CHINESE MOUNTAIN CAT
Felis bieti

Plate 1

CHINESE DESERT CAT, CHINESE STEPPE CAT

HB 68.5–84cm; T 32–35cm; W 6.5–9kg

Light yellow-grey in winter, darkening to tawny or grey-brown in summer, with very faint markings or none at all except for a dark dorsal line. Tail bushy, conspicuously banded, with a dark tip. Ears tufted. Limited genetic data suggest this is a subspecies of African Wildcat, but classification is still disputed. Distribution and Habitat Known only from Qinghai, Sichuan and Gansu provinces, C China. Records from elsewhere in China and Tibet are equivocal. Inhabits alpine grassland, meadows, shrubland and forest edges at 2,500–5,000m. May also occur in true montane forest, semi-desert and cold desert, but this is unconfirmed. Feeding Ecology Virtually unstudied in the wild. A single study on diet showed that small mammals such as voles, mole rats, hamsters and pikas make up 90% of the diet. Also eats birds, including records of pigeons, partridges and pheasants. Social and Spatial Behaviour Unknown. Probably solitary. Dens in rock outcrops, burrows under tree roots and dense thickets. Reproduction and Demography A handful of records suggest seasonal breeding, which is likely given C China’s harsh winters. Male–female pairs are mostly observed January–March, which is the likely breeding season, with kittens born around May. MORTALITY and LIFESPAN Unknown. Status and Threats Very restricted distribution, and thought to be naturally rare. Killed for its fur, which is mostly for local use, but hunting is widespread and pelts are common in fur markets. Large-scale government-mandated poisoning of rodents and lagomorphs prevalent in C China is likely to constitute a serious threat both by reducing prey populations and secondary poisoning. CITES Appendix II; Red List VU, population trend Decreasing.

EUROPEAN WILDCAT
Felis silvestris,
AFRICAN WILDCAT
Felis lybica

Plate 1

HB 40.6–64cm, 44–75cm; T 21.5–37.5cm; W 2–5.8kg, 2–7.7kg

Considered conspecific until 2017, now classified as 2 distinct species that diverged ~1 million years ago. The domestic cat first arose from the African Wildcat in the Fertile Crescent more than 9,000 years ago, with later genetic contributions in Egypt, and possibly also from the European Wildcat in SE Europe. Domestic cats hybridise with both wildcats and the wild species likely interbreed where distributions abut around the Caspian Sea. Wildcats closely resemble the domestic cat, although wild individuals are generally larger, taller and more robust. African Wildcat: in Africa/Middle East, sandy grey to pale beige with banded legs and red-backed ears; in Asia (‘Asiatic Wildcat’, possibly a distinct subspecies, F. l. ornata), typically spotted on an isabelline background with noticeable ear-tufts. European Wildcat looks like a heavily built striped tabby with a bushy tail, white chin and white throat. Piebald, ginger and black variants are usually the result of hybridisation with domestic cats. Distribution and Habitat European Wildcat: W Europe from N Scotland to the W Caspian. African Wildcat: Africa (except C Africa and the Sahara), the Middle East, and much of W and S Asia to C China. Very broad habitat tolerance, from sea-level to 3,000m. European Wildcats inhabit mainly temperate forest, woodland, scrub and associated habitats. African Wildcats occur in virtually all habitats with cover, excluding dense forest and open desert interiors. Both species avoid very open habitat, including high montane areas and deep snow. Both readily occupy agricultural lands, fields with cover and plantations, but avoid intensively farmed habitat with little cover. Feeding Ecology Through almost their entire ranges, diet of both species is dominated by small rodents such as mice, rats, voles, jirds, gerbils and jerboas; a notable exception is European Wildcat in Scotland, where European Hares and European Rabbits comprise up to 70% of prey. Other important prey includes small birds, especially ground foragers such as doves, pigeons, partridges, sandgrouse, guinea fowl, quails and weavers. Reptiles, including large venomous snakes (e.g. Puff Adders and cobras), amphibians and arthropods are also eaten. Readily kills poultry and (rarely) very young domestic goats and lambs. Drinks daily when water is available; African Wildcat often occurs far from water in the Kalahari, Namib and Sahara, suggesting it is water-independent. Hunts mostly on the ground, and is chiefly nocturno-crepuscular; when protected, may be active during the day, especially in cold winters. Scavenges, and sometimes caches food by covering it with debris, soil and leaf litter. Social and Spatial Behaviour Solitary and territorial. Displays typically feline territorial behaviours such as marking with urine and faeces, but extent of territorial defence probably varies widely between different habitats. Range size also varies widely, typically (but not universally) with larger male ranges overlapping multiple female ranges. Range estimates include 1.7–2.75km2 (s) to 13.7km2 (1 ; Portugal), 1.75km2 (average for both sexes; E Scotland with abundant lagomorphs), 3.5km2 (s, average) to 7.7km2 (s, average; S Kalahari), 8–10km2 (both sexes; W Scotland with scarce prey), 11.7km2 (average for both sexes; Saudi Arabia) and 51.2km2 (1 ; United Arab Emirates). Density 0.7–10/km2. Reproduction and Demography Breeds seasonally in areas with extreme seasonality, i.e. most of European Wildcat range, and the Sahara, mating winter–early spring and giving birth spring–early summer. Elsewhere, kittens may be born year-round, although birth peaks often coincide with prey flushes during or after the rainy season, e.g. E and South Africa. Gestation 56–68 days. Litter size typically 2–4, rarely to 8. Weaning at 3–4 months, independent at 5–10 months. Sexually mature at 9–12 months, but wild individuals probably first breed at 18–22 months. MORTALITY Poorly documented; most mortality in studied populations is due to human factors. Known predators include large cats, Golden Eagle, Honey Badger (of kittens) and domestic dogs. Approximately 20 Meerkats killed a 6-week-old African Wildcat kitten after pulling it from a burrow in the Kalahari, South Africa. Starvation of kittens and subadults contributes to low survival in harsh winters. LIFESPAN 11 years in the wild, 15 in captivity. Status and Threats Both species are widely distributed, adaptable and tolerant of human activity such as agriculture and forestry, which often elevates rodent populations. Not endangered in any traditional sense, but both hybridise with domestic cats (producing fertile hybrids), especially in W Europe; in Scotland, up to 88% of wild-living European Wildcats may be hybrids. Only remote populations are thought to entirely lack hybrids, although hybridisation in many rural areas is not as extensive as once assumed. Other important threats include persecution for killing poultry and for perceived killing of game species (Europe) and small stock, hunting for fur (mainly C Asia), domestic cat diseases and roadkills. Both species: CITES Appendix II; Red List LC, population trend Decreasing.

Plate 1

BLACK-FOOTED CAT
Felis nigripes

Plate 2

SMALL-SPOTTED CAT

HB 35.3–41.5cm, 36.7–52cm; T 12–20cm; W 1–1.6kg, 1.5–2.58kg

One of the smallest cats. Light tawny to buff-grey with ginger to black markings. Northern individuals are generally paler, with the darkest, most strongly marked animals found in Eastern Cape, South Africa, but there is wide variation within populations. Distribution and Habitat Endemic to southern Africa and restricted to open, short grass habitat, dry savannah, Karoo scrub and semi-desert. Independent of drinking water, but does not occur in open hyper-arid desert interiors. Feeding Ecology Specialises in rodents <100g and ground-roosting birds to the size of Black Bustard. Heaviest prey is Cape Hare; attacks on newborn Springbok lambs are unsuccessful. Reptiles, amphibians, eggs and invertebrates are also eaten. Nocturno-crepuscular, hunting up to 70% of the night with 3 strategies: rapid bounding through cover to flush birds; painstaking weaving around cover searching for prey; and waiting in ambush at rodent burrows. Succeeds in 60% of hunts (Benfontein NR, South Africa), with 10–14 rodents or birds caught per night, averaging a kill every 50 minutes. Surplus food is cached in shallow diggings or hollow termitaria; scavenges. Social and Spatial Behaviour Solitary and territorial, with very frequent urine-marking and vocalising. Male ranges encompass up to 4 female ranges. Males guard oestrous females and fight intruding males. Ranges average 8.6km2 (s) and 16.1km2 (s). Density estimates up to 16.7/100km2 (high-quality habitat; typically at much lower densities). Reproduction and Demography Seasonal (South Africa). Birth peaks coincide with spring–summer rains and prey flushes. Oestrus 36 hours; gestation 63–68 days. Litter size 1–4. Kittens weaned by 2 months and independent at 3–4 months. Sexual maturity at 7 months (s) and 9 months (s). MORTALITY Poorly known; predators include Black-backed Jackal, Caracal, domestic dogs and large owls. LIFESPAN 8 years in the wild, 16 in captivity. Status and Threats Probably naturally uncommon. Threatened by agricultural expansion into semiarid areas and associated overgrazing, burning and insecticides, which impact rodent and insect populations. Hundreds of Black-footed Cats are killed annually during control activities intended for jackals in South Africa. CITES Appendix I; Red List VU, population trend Decreasing.

SAND CAT
Felis margarita

Plate 2

HB 39–52cm, 42–57cm; T 23.2–31cm; W 1.35–3.1kg, 2–3.4kg

Very small, strikingly pale cat with indistinct markings on the body, resolving to dark stripes on the legs and tail. W African cats are smaller, paler and more strongly marked than eastern individuals, which are typically darker grey in tone. Flat, broad head topped by oversized ears. Feet densely covered in dark fur, probably for traction and insulation on loose hot sand. Distribution and Habitat N Africa, the Middle East and C Asia. A desert specialist, capable of occupying true desert with rainfall of <20mm/year. Inhabits a variety of sandy and stony desert habitats with cover, and arid shrub-covered steppes. Absent from heavily vegetated valleys in these habitats. Feeding Ecology Eats mainly small rodents, including gerbils, spiny mice, jirds and jerboas, plus occasional kills of young hares, small birds, reptiles (including venomous snakes) and invertebrates. Independent of drinking water. Foraging is mainly nocturnal. Capable of very rapid digging to excavate burrowing prey, and sometimes caches food with a covering of sand. Social and Spatial Behaviour Solitary. Poorly known but preliminary radio-tracking data from Morocco indicates exceptionally large home ranges likely to average ~200km2 for both sexes; over 9–10 months, 1 female covered 760km2 and 3 males covered 150km2 to >500km2. A fourth male, possibly a disperser, covered 1,500km2 over 9 months. Nightly straight-line movements average 5.4km, up to 21km. Reproduction and Demography Breeding appears seasonal in the Sahara. Mating November–February; births January–April. Oestrus 5–6 days; gestation 59–67 days. Litter size 1–6, rarely >4, average 2.7 (captivity). Weaning at around 5 weeks, and independence from 4 months. Sexually mature at 7–14 months (captivity). MORTALITY Unknown, but likely vulnerable to African Wolf and large raptors. LIFESPAN 16 () to 19 () years in captivity, significantly less in the wild. Status and Threats Probably naturally rare, but its habitat is so remote that it is somewhat insulated from human activities. Threats include expansion of cultivation, and feral domestic cats and dogs, which result in predation, competition and possible disease transmission. Killed around human settlements in traps for canids. CITES Appendix II; Red List LC, population trend Unknown.

JUNGLE CAT
Felis chaus

Plate 2

SWAMP CAT, REED CAT

HB 56–85cm, 65–94cm; T 20–31cm; W 2.6–9kg, 5–12.2kg

Leggy cat with a short tail. Uniformly coloured, with indistinct body markings that become more obvious on the limbs. Temperate animals tend to be darker and more richly marked than tropical individuals. Melanism reported from India and Pakistan. Distribution and Habitat Temperate and tropical S Asia, extending into Egypt along the Nile Valley. Prefers dense reed-beds, long grass and scrub habitats in swamps, and wetland and coastal areas, but also inhabits dry and evergreen forests. Tolerates cultivated marshy landscapes, including sugar-cane fields and rice paddies. Feeding Ecology Small mammals (<1kg) comprise primary prey, especially small rodents, Muskrat, squirrels and hares; Coypu (5–9kg), and neonates of gazelle and Chital occasionally recorded. Birds make up second-most important prey category, especially waterfowl, francolins, pheasants, peafowl and jungle fowl. Also eats small reptiles and amphibians, and has been observed diving into shallow water for fish. Recorded eating large quantities of Russian Silverberry fruits during winter (Tajikistan). Readily preys on domestic poultry, and sometimes scavenges from the kills of larger carnivores and people’s furbearer traps. Social and Spatial Behaviour Poorly known. Solitary. Typical feline scent-marking and vocalisation suggest maintenance of exclusive core areas. Larger male home ranges likely overlap several smaller female ranges. There are no density estimates, but it is frequently the most common felid where it is found. Reproduction and Demography Thought to be weakly seasonal. Most observed mating November–February. Kittens born December–June. Gestation 63–66 days. Litters average 2–3, exceptionally to 6. Kittens independent at 8–9 months. Sexually mature at 11 months (s) and 12–18 months (s). MORTALITY Unknown. LIFESPAN 20 years in captivity. Domestic dogs are a confirmed predator in India. Status and Threats Common in many parts of its range and tolerates agricultural landscapes with cover. Destruction and development of wetland habitats are a particular threat in arid areas, e.g. Egypt. Persecuted for poultry raiding, trapped heavily around Coypu fur farms (former USSR) and hunted for fur in its northern range. CITES Appendix II; Red List LC, population trend Decreasing.

Plate 2

PALLAS’S CAT
Otocolobus manul

Plate 3

MANUL, STEPPE CAT

HB 46–53cm, 54–57cm; T 23–29cm; W 2.5–5kg, 3.3–5.3kg

Stocky, heavily furred small cat, silvery grey to rufous-grey with faint striping on the body. Long winter coat has a pale, frosted appearance; spring–summer coat is darker, with more obvious stripes and often a reddish tinge. Face has dark cheek stripes, and the crown is distinctively marked with small spots. Bushy tail is banded with narrow stripes and ends in a dark tip. Coloration provides excellent camouflage in open rocky habitat: Pallas’s Cat is poorly adapted for running, and when threatened freezes and flattens itself to the ground, conferring very effective concealment. Distribution and Habitat C Asia, from the Caspian Sea through N Iran, Afghanistan, Pakistan and N India to C China, Mongolia and S Russia. Confirmed by camera-trap from Bhutan and Nepal in 2012. Lives at 450–5,073m in cold, arid habitats with cover, especially dry grassland steppes with stone outcrops and stony semi-desert. Prefers valleys and rocky areas, and avoids completely open habitat. Although well adapted for extreme cold, usually avoids areas with deep snow. Feeding Ecology Hunts mainly small rodents and lagomorphs. Pikas are especially important prey, typically comprising more than 50% of the diet; voles, mice, hamsters, gerbils, ground squirrels and marmots are also important. Occasional prey includes hares, hedgehogs, small birds, lizards and invertebrates. There is 1 verified record of predation on newborn Argali lamb (Mongolia). Hunts by 3 distinct techniques: ‘stalking’, by carefully creeping around cover; ‘moving and flushing’, used mainly in long summer undergrowth; and waiting in ambush at rodent burrows. Not known to kill any livestock or poultry. Recorded scavenging from carcasses. Social and Spatial Behaviour Solitary. Both sexes maintain enduring home ranges, with large overlapping male ranges encompassing multiple smaller female ranges that overlap minimally with each other. Likely to be territorial, at least in the breeding season; breeding males often have injuries consistent with fighting. Female territories 7.4–125.2km2, averaging 23.1km2, compared with male territories of 21–207km2, averaging 98.8km2 (Hustain Nuruu NP, Mongolia). Density estimates 2–8/100km2. Reproduction and Demography Highly seasonal. Mating December–March; births late March–May. Oestrus very short at 24–48 hours; gestation 66–75 days. Litters average 3–4, exceptionally to 8 (captivity). Kittens independent at 4–5 months. Sexual maturity at 9–10 months for both sexes. MORTALITY 68% of kittens do not survive to disperse, 50% of adults (>1 year old) survive to age 3 (Hustain Nuruu NP). Most mortality occurs in winter in October–April. Known predators include large eagles, Red Fox and domestic dogs. LIFESPAN 11.5 years in captivity. Status and Threats Lives in remote areas but at low densities. Poorly adapted to avoid predators, and depends on fairly specific habitats, making it naturally vulnerable to threats. Hunted for fur in much of its range, and domestic dogs often constitute a key predator; human factors (including dogs) account for 56% of deaths in C Mongolia. State-sanctioned rodent-poisoning campaigns in China, Mongolia and Russia are a serious threat to Pallas’s Cat prey. Captives, especially kittens, are very vulnerable to toxoplasmosis (assumed contracted in captivity), which is often fatal. Legally hunted without limits in Mongolia. CITES Appendix II; Red List NT, population trend Decreasing.

FISHING CAT
Prionailurus viverrinus

Plate 3

HB 57–74.3cm, 66–115cm; T 24–40cm; W 5.1–6.8kg, 8.5–16kg

Robust cat with a powerful, blocky head and short, thick tail. Olive-grey fur covered in dark spots that often coalesce into stripes on the nape, shoulders and back. Feet partially webbed, with large claws that are protractile but protrude partially from the claw-sheaths. Fur is not waterproof, as sometimes claimed. Distribution and Habitat Southeast Asia from NE Bangladesh and NE India to S Thailand, and isolated populations in Nepal, extreme SE Pakistan (confirmed Chotiari wetlands, Sindh province, 2015), SW India and Sri Lanka. No confirmed records from Java since 2000, and its presence in Sumatra has never been established. Closely associated with wetland habitats, including marshes, riverine woodland, dense Terai grassland (Nepal) and mangroves. Sometimes found in degraded habitats around aquaculture ponds and rice paddies, but generally intolerant of wetland modification. Feeding Ecology Paws and claws are adapted for aquatic foraging and a diet dominated by fish. Capable swimmer that submerges itself in pursuit of prey, and hunts along the water’s edge or in shallows, where it scoops up fish with its paws. Despite its aquatic adaptations, its dentition is more generalised, indicating a broader diet. Also kills small mammals, birds (including ducks and coots hunted in the water), reptiles, amphibians and invertebrates. Occasionally kills prey to the size of Chital fawns. Sometimes kills livestock, mainly very young goats and poultry; there are credible but unconfirmed records of kills of neonate calves. Reports of killing children are extremely unlikely. Scavenges. Social and Spatial Behaviour Poorly known. Solitary. Range sizes 2–6.8km2 (4 s) and 4–13.5km2 (2 s; mostly agri-aquacultural habitat, Thailand) and 4–6km2 (2 s) and 22km2 (1 ; Terai grassland, Chitwan NP, Nepal). Scats are apparently deposited in latrines. Reproduction and Demography Unknown from the wild. In captivity, gestation 63–70 days, litter size 1–4. Sexual maturity in 1 captive female occurred at 15 months. MORTALITY Poorly known; of 11 known deaths during a Thai study, 5 were caused by people, the rest to unknown causes. LIFESPAN 12 years in captivity. Status and Threats Until recently, considered widespread and relatively common, but accelerated development of wetlands and floodplains throughout Asia, especially for aquaculture, is a pervasive threat and a key factor in rapid and widespread declines. Illegal persecution, mostly as a perceived threat to poultry and aquaculture, exacerbates declines. Now rare in Java, Laos and Vietnam, and has lost significant range in India and Thailand. May no longer occur in Pakistan. Main strongholds appear to be S Thailand, Sri Lanka and isolated areas of Bangladesh, NE India and Nepal. CITES Appendix II; Red List VU, population trend Decreasing.

Plate 3

MAINLAND LEOPARD CAT
Prionailurus bengalensis
SUNDA LEOPARD CAT
Prionailurus javanensis

Plate 4

INDOCHINESE LEOPARD CAT (P. bengalensis)

HB 38.8–65.5cm, 43–75cm; T 17.2–31.5cm; W 0.55–4.5kg, 0.74–7.1kg

Considered conspecific until 2017, now classified as 2 species that diverged an estimated >900,000 years ago and are morphologically and genetically distinct. A hybrid zone occurs on the Thai–Malay peninsula, provisionally classified as P. bengalensis, with genetic and morphological characteristics of both. Preliminary molecular analysis suggests 2 Sunda subspecies, P. j. javanensis in Java and Bali, and P. j. sumatranus in Sumatra, Borneo and the Philippines; and 2 highly variable subspecies on continental Asia, divided into a southern subspecies, P. b. bengalensis, occurring from Afghanistan and India to approximately E China, and a northern subspecies, the Amur Leopard Cat (P. b. euptilurus), of the Russian Far East, NE China, the Korean Peninsula, Taiwan, and Japan’s Tsushima and Iriomotejima islands. Sunda Leopard Cats are the smaller and generally less variably coloured and patterned of the 2 species, typically with small, solid dark spots and dabs on a muted background colour that varies widely from drab ginger-brown to very dark brown. Mainland Leopard Cats vary along a south–north cline; southernmost individuals are small and richly coloured, becoming larger and generally plainer in the north. Individuals in mainland tropical Asia have rich yellow to tawny-brown or ginger-brown fur, and bold markings varying from large solid spots and rosettes to blotches with dark tawny or reddish centres. Amur Leopard Cats in temperate Russia, Korean Peninsula and China are very pale ginger-grey to silver-grey in winter, with long, dense fur that moults to a darker summer coat of russet-brown to dark grey-brown. Complete melanism has not been recorded in either species, although there are occasional records of pseudomelanistic individuals with extensive enlarging and coalescing of the dark markings.

Distribution and Habitat

Sunda Leopard Cat: Sumatra, Java, Bali, Borneo, and the Philippine islands of Palawan, Panay, Negros and Cebu. Possibly introduced by humans to some or all of its Philippines’ range. It may occur on the Malay Peninsula south of the Kra Isthmus (currently regarded as P. bengalensis). Mainland Leopard Cat: very widely distributed in tropical and temperate mainland Asia, from the Russian Far East, China, Taiwan, Japan (Tsushima and Iriomotejima islands) throughout Southeast Asia to S India, N Pakistan, E Afghanistan and extreme S Tajikistan. Both species inhabit all forest types, woodland, scrub, shrublands, marshes, wetlands and mangroves, from sea-level to at least 3,000m (4,250m recorded for Mainland Leopard Cat, Arunachal Pradesh, India). Amur Leopard Cats inhabit vegetated valleys in cold temperate forest with winter snowfall, limited to areas with shallow snow. Both species largely avoid open grasslands, steppes and rocky areas lacking vegetation. Leopard cats inhabit human-modified habitats provided there is cover, including in secondary forest, sugar-cane fields, rice paddies and plantations of oil palm, coffee, rubber and tea. They occur very close to human habitation, including in suitable habitat patches in major metropolises, e.g. Miyun Reservoir and Yeyahu NR, Beijing.

Feeding Ecology

Leopard cats hunt a wide variety of small vertebrates, chiefly small rodents (especially mice, rats and squirrels), tree shrews, shrews, birds, snakes and lizards. Both species are surprisingly aquatic, and forage in shallow water for freshwater crabs, amphibians and invertebrates. Records of larger prey, e.g. langurs, hares, Lesser Mouse Deer and Wild Boar, were probably scavenged. Amur Leopard Cats in Russia reputedly kill unguarded neonate (<1-week-old) ungulates, including Siberian Roe Deer, Sika Deer and Long-tailed Goral. Introduced Black Rats, skinks, snakes, frogs and a large cricket species are the main prey of Iriomotejima cats. Harmless to hoofstock, but they take domestic poultry and are easily baited with chickens. Hunting activity is variable, ranging from strictly nocturnal at some sites to cathemeral.

Social and Spatial Behaviour

Solitary and apparently weakly territorial. Male ranges generally overlap numerous smaller female ranges (although range size differs little between sexes in some populations, e.g. Phu Khieo WS, Thailand). Overlap between same-sex adults is considerable at range edges, and usually minimal in exclusive core areas (which overlap significantly in Phu Khieo). Range size 1.4–37.1km2 (s) and 2.8–28.9km2 (s). Leopard cats are often the most abundant felid in most of the respective range. Density estimates 17−22/100km2 (subtropical–temperate Himalayan forest, Khangchendzonga BR, India), 34/100km2 (Iriomotejima), 37.5/100km2 (Tabin WR, Sabah).

Reproduction and Demography

Breeding varies from highly seasonal in Russia to aseasonal in the tropics; Sunda Leopard Cats are assumed to be entirely aseasonal. Captive individuals are able to have 2 litters a year, although a single litter is probably typical in the wild. Gestation 60–70 days. Litter size 1–4. Sexual maturity at 8–12 months (captivity). MORTALITY Estimated annual adult mortality varies from 8% (remote sanctuary, Phu Khieo) to 47% (accessible protected area, Khao Yai NP, Thailand). Probably vulnerable to a wide variety of predators; records confirmed for Leopards, Reticulated Pythons and domestic dogs. Humans are the main source of mortality in many areas, mainly from hunting, persecution and roadkills (especially on Iriomotejima and Tsushima). LIFESPAN 13 years in captivity.

Status and Threats

Both species are widespread, adaptable and reach high densities in suitable natural and anthropogenic habitats. The Mainland Leopard Cat is legally killed for fur and is heavily hunted in its temperate range, especially in China, where densities are naturally low. High fur harvests are likely to produce declines. In tropical and subtropical Asia, both species are widely killed for fur and meat, in retaliation for poultry predation, and are targeted for the pet trade. Many island populations are small and threatened by rapid development; populations are declining on Tsushima (Mainland Leopard Cat), and Panay, Negros and Cebu (Sunda Leopard Cat). Mainland Leopard Cat: CITES Appendix I – Bangladesh, India, Thailand, Appendix II – elsewhere; Red List LC (CR on Iriomotejima), population trend Stable. Sunda Leopard Cat: CITES Appendix II; Red List LC (VU in the Philippines), population trend Stable.

Plate 4

RUSTY-SPOTTED CAT
Prionailurus rubiginosus

Plate 5

HB 35–48cm; T 15–29.8cm; W 1–1.1kg, 1.5–1.6kg

One of the smallest cats, the size of a slight, very small domestic cat. Rufous-brown or grey-brown with rows of reddish to dark brown spots that sometimes form complete stripes on the nape, shoulders and upper flanks. White or pale cream underparts. Tail solid rust-brown, sometimes with faint bands. Distribution and Habitat Endemic to India, Sri Lanka and SW Nepal (confirmed by photographs from Bardia NP, 2012, and Suklaphanta NP, 2016). Putative range abuts extreme SE and NE Pakistan, where it may occur marginally. Once regarded as a moist forest specialist, now revealed through camera-trapping to have a much wider distribution and habitat range, including dry forest, bamboo forest, wooded grassland, arid shrubland, scrubland and vegetated rocky habitats. Recorded to 2,100m. Occurs in modified habitats, including cropland such as maize and rice (which harbour abundant rodents and amphibians), tea plantations and occasionally in abandoned dwellings in villages. Feeding Ecology Poorly known. Has a reputation for being especially fierce and taking outsized prey, based on observations in unnatural settings; an 8-month-old individual in captivity almost killed a tame gazelle lamb several times its size by a throttling throat bite before people intervened; such attacks in the wild are highly unlikely. The known diet is mainly small vertebrates such as shrews, Indian Gerbil, bandicoot rats and mice, especially the Little Indian Field Mouse; small birds, hatchlings, reptiles, amphibians and invertebrates are also eaten. It sometimes kills free-ranging domestic poultry, mainly chicks; it rarely, if ever, enters coops or dwellings for poultry. Most sightings of foraging cats are on the ground, but it is a highly agile climber and possibly hunts both on the ground and arboreally. An adult caught a shrew after a 50m ground chase that began while the cat was apparently foraging in the low branches of a tree. Chiefly nocturnal. Social and Spatial Behaviour Unknown. Reproduction and Demography Unknown from the wild. In captivity, reproduction is aseasonal; 2 wild litters, 1 each from India and Sri Lanka, both found in February. Gestation 66–79 days, usually 67−71 days. Litter size 1–3. MORTALITY Unknown but likely vulnerable to predation by many larger carnivores; a mother with 2 young kittens was observed losing one to an Indian Cobra (Sri Lanka). LIFESPAN 12 years in captivity. Status and Threats Regarded as rare. Recent observations have dramatically expanded the known range, although it is not considered common anywhere. Found fairly often in association with human settlements; given its tiny size and potential utility in controlling rodents, it can prosper provided it is tolerated by people. However, it is often killed for skins and meat, by domestic dogs and as a perceived threat to poultry. Unintentional mortality likely results from very widespread use of insecticides and rodenticides in rural S Asia. The cats are sometimes mistakenly persecuted as Leopard cubs (Sri Lanka). CITES Appendix I – India, Appendix II – elsewhere; Red List NT, population trend Decreasing.

FLAT-HEADED CAT
Prionailurus planiceps

Plate 5

HB 44.6–52.1cm, 41–61cm; T 12.8–16.9cm; W 1.5–1.9kg, 1.5–2.2kg

Very small, unusual cat with a short tubular body; stubby tail; compact, foreshortened face with a flattened forehead; large, closely set eyes; and small ears. Feet partially webbed and claws protrude partially from reduced claw-sheaths; the claws are protractile. Body is dark roan-brown, becoming rusty brown on the head, with white to whitish-grey underparts. Face is bright white on muzzle, cheeks and under the eyes, with contrasting dark rusty-brown forehead, cheek and eyebrow stripes. Body is largely unmarked except for light dappling and banding on the underparts, chest and lower limbs; the tail is sometimes faintly banded. Distribution and Habitat Peninsular Malaysia (possibly also just across the border into S Thailand), Borneo and Sumatra. Very strongly associated with low-elevation moist forested habitats; >70% of records are within 3km of large rivers or water sources, and >80% occur below 100m. Inhabits primary and secondary forest, peat-swamp forest, mangrove and coastal scrub forest. Reports from oil palm and rubber plantations suggest some tolerance of habitat modification, but records from such habitats are dubious or very uncommon. Feeding Ecology Very poorly known, but the unique morphology, behaviour and habitat preferences of the species suggest specialisation for hunting aquatic prey in shallow water and along muddy riverbanks. Captive animals are attracted to water, readily submerging themselves and feeling for food in pools with spread paws, similar to raccoons. Stomach contents of dead wild individuals contained fish and crustaceans. Captives adeptly kill rodents with a typically felid nape bite; wild individuals almost certainly take small mammals, reptiles and amphibians. They are sometimes killed in traps set at poultry coops, suggesting they take domestic fowl. Chiefly nocturnal based on camera-trap images. Social and Spatial Behaviour Flat-headed Cats have never been radio-collared and socio-spatial behaviour is unknown. Always the least common felid in camera-trap surveys, even when targeted, suggesting naturally very low densities. Reproduction and Demography Unknown from the wild. Gestation 56 days (captivity). Litter size 1–2 (based on only 3 captive births). MORTALITY Unknown. Presumably vulnerable to a wide array of predators; occasionally killed by people and dogs, and on roads. LIFESPAN 14 years in captivity. Status and Threats Known from ~110 physical records and sightings; despite extensive camera-trapping efforts, 44 of 46 known photos come from just 4 sites in Borneo, with a single image each from Peninsular Malaysia and Sumatra. The species’ relatively restricted distribution and very close association with moist, forested habitats makes it particularly vulnerable to habitat loss. As of 2009, an estimated 54–68% of suitable Flat-headed Cat habitat had been converted by people, especially clearing and draining of forest wetlands for croplands and forestry. Overfishing and freshwater pollution from agriculture and mining are likely to exacerbate declines driven by habitat loss, and hunting by people may exert strong local effects; Flat-headed Cat skins often occur in longhouses in Sarawak. Live animals (usually kittens) occasionally appear in the pet trade. Many authorities now consider the Flat-headed Cat to be Southeast Asia’s most threatened small felid. CITES Appendix I; Red List EN, population trend Decreasing.

Plate 5

MARBLED CAT
Pardofelis marmorata

Plate 6

HB 45–62cm; T 35.6–53.5cm; W 2.5–5kg

Resembles a small clouded leopard, with thick grey-buff to red-brown fur, patterned with large, dark-bordered blotches that become small dabs on the limbs. Tubular bushy tail proportionally very long, sometimes exceeding the head–body length and distinctive in the field. During relaxed walking, the tail is held horizontally in a continuous straight line from the body. Melanism occurs very rarely. Distribution and Habitat Southeast Asia, south of the Himalayas in Sikkim, India, to SW China, and through Indochina, Borneo and Sumatra. It may occur in E Nepal, although there are no unequivocal records. Restricted to forested habitats, chiefly undisturbed evergreen, deciduous and tropical forests. Can occupy secondary and logged forests, although it is unknown whether modified habitat is suboptimal. Feeding Ecology Except for 1 radio-collared female tracked for a month in Thailand, the species has never been studied by telemetry in the wild. Diet is likely to be dominated by small vertebrates. Highly agile climber and has been observed hunting in trees, perhaps for arboreal mammals such as squirrels, as well as birds. From limited camera-trapping in protected areas, thought to be mostly diurnal. Social and Spatial Behaviour Virtually unknown. Occasional sightings of adult pairs have fostered speculation that it forms long-term pair bonds, but it is more likely to be solitary. A collared Thai female used a range of 5.3km2 in 1 month. Density estimates 5/100km2 (upland forest, Dampa TR, India), 7.1/100km2 (upland forest, Tawau Hills Park, Borneo), 8.8/100km2 (upland forest, Htamanthi WS, Myanmar), 10.45/100km2 (selectively logged lowland forest, Tabin WR, Borneo), 19.6/100km2 (protected lowland forest, Danum Valley CA, Borneo). Reproduction and Demography Very poorly known. Gestation 66–82 days (captivity). Litters average 2 kittens (based on only 2 captive births). Females sexually mature at 21–22 months (captivity). MORTALITY Unknown. LIFESPAN 12 years in captivity. Status and Threats Reaches high densities in protected forest but seems highly forest-dependent and is very vulnerable to habitat loss. Hunting is very prevalent throughout its range and likely exacerbates declines in concert with deforestation. CITES Appendix I; Red List NT, population trend Decreasing.

BAY CAT
Catopuma badia

Plate 6

HB 53.3–67cm; T 32–39.1cm; W (1 emaciated ) 2kg

Resembles a small, slender Asiatic Golden Cat with a proportionally smaller, rounded head and stubby, rounded ears. Occurs in 2 morphs: rich, rusty red; and grey with variable red undertones, especially along the transition from the upper body colour to the paler underparts. Unmarked except for stripes on the forehead and cheeks, and faint spotting along the transition between the upper body colour and pale underparts. Bright white underside to the tail with a dark dorsal tip that is distinctive in the field. Distribution and Habitat Endemic to Borneo. Closely associated with densely forested habitats, with most historical records in primary, riverine, swamp and mangrove forests. Tolerates moist plantation forests with a dense understorey, and has been camera-trapped from recently logged secondary forest, suggesting some tolerance for habitat modification. Feeding Ecology Unknown, but presumably small vertebrates make up major food items. Two Bay Cats were trapped in 2003 when they entered an animal dealer’s pheasant aviaries, suggesting that the species may attack domestic poultry. Social and Spatial Behaviour Unknown. Rarely photographed during camera-trapping surveys, suggesting that it occurs at very low densities, e.g. Bay Cats were photographed 25 times at 4 sites in eastern Sabah over 4 years, compared with 259 images of Sunda Clouded Leopards and more than 1,000 images of Sunda Leopard Cats. Reproduction and Demography Unknown. Status and Threats Apparent extreme rarity of the species and its dependence on forest raise concerns for its conservation prospects. Forest conversion, especially to oil palm plantations, is regarded as a serious threat. The species’ rarity and value are known to animal dealers, elevating illegal trapping pressure. CITES Appendix II; Red List EN, population trend Decreasing.

ASIATIC GOLDEN CAT
Catopuma temminckii

Plate 6

TEMMINCKS GOLDEN CAT

HB 66–94cm, 75–105cm; T 42.5–58cm; W 8.5kg, 12–15.8kg

Usually rich russet-brown, but varying from pale tawny to dark greyish brown. Largely unmarked except for the face and faint spotting on the chest and belly. A richly spotted ‘ocelot’ morph is recorded from Bhutan, China and Myanmar. Melanism occurs. Except in black individuals, the underside of the tail is always conspicuously bright white with a dark upper tip. Distribution and Habitat Sub-Himalayan Nepal, NE India and Bhutan to S China, Southeast Asia and Sumatra. Found in a variety of moist and dry forests, usually below 3,000m, but higher on Himalayan slopes, e.g. 3,738m (Bhutan) to 4,369m (Myanmar). Has been sighted or killed near human settlements, including in open agricultural areas, and appears to be more tolerant of open habitat than clouded leopards, Marbled Cat and Bay Cat; even so, it is never far from cover. Feeding Ecology Poorly known. Confirmed prey includes mice, rats, Berdmore’s Ground Squirrel, mouse deer, Dusky Leaf Monkey (~6.5kg), snakes, lizards and birds. Powerfully built and reputed to kill medium-sized ungulates, including muntjacs and livestock to the size of neonate cattle and buffalo calves (which is unlikely). Confirmed records of livestock kills are mostly from hunters shooting the cat over depredated carcasses, in which it may have been the predator or possibly only a scavenger. Sometimes raids poultry. Nocturno-crepuscular, but diurnal activity is recorded under protection. Social and Spatial Behaviour Poorly known. Solitary. Only range sizes known are 32.6km2 (1 ) and 47.7km2 (1 ) from Phu Khieo WS, Thailand. Reproduction and Demography Unknown from the wild. In captivity, reproduction aseasonal, gestation 78–80 days and litter size 1–3 (typically 1). Sexual maturity in captive animals at 18–24 months. MORTALITY Unknown. LIFESPAN 17 years in captivity. Status and Threats Threatened by forest loss and illegal hunting, which are widespread throughout its range, but status and degree of threat are poorly known. Skins of Asiatic Golden Cats are traded heavily in China and Myanmar, where hunting pressure is regarded as high. CITES Appendix I; Red List NT, population trend Decreasing.

Plate 6

SERVAL
Leptailurus serval

Plate 7

HB 63–82cm, 59–92cm; T 20–38cm; W 6–12.5kg, 7.9–18kg

Dappled with bold spots, coalescing into blotches on the extremities and nape. A buff-coloured morph with a faint freckled appearance (‘Servaline’) occurs in W and C Africa. Melanism is recorded chiefly in Equatorial highlands and forest-savanna ecotone. Distribution and Habitat Africa. Inhabits mainly woodland savannah, grassland, forest and alpine moorland to 3,850m, usually associated with rivers, marshes and floodplains. Absent from rainforest and desert. Tolerates agricultural areas with cover. Feeding Ecology Specialises in hunting small mammals in long grass or shrubby habitats, with rodents and shrews comprising 80–93.5% of the diet. Small grassland birds are the next most important prey, and it sometimes hunts large birds, including flamingos and storks. Ancillary prey includes genets, mongooses, hares, juvenile antelopes, small reptiles and arthropods. Livestock depredation is unusual, but it occasionally kills poultry and untended young goats. Hunting is mostly crepuscular, becoming nocturnal near humans. Rarely scavenges. Social and Spatial Behaviour Solitary and territorial, although same-sex adults appear relatively tolerant. Home ranges overlap considerably. Range size 15.8–19.8km2 (s) and 31.5km2 (1 ) in KwaZulu-Natal, South Africa. Density estimates 9.9/100km2 (Luambe NP, Zambia, with full carnivore complement) and 41/100km2 (optimum protected habitat, Ngorongoro Crater, Tanzania), to 76/100km2 (Secunda CTL fuel plant, South Africa, a fenced industrial complex with extensive moist grassland, very high rodent abundance and no other carnivores). Reproduction and Demography Breeding appears weakly seasonal; births peak November–March (southern Africa) and August–November (Ngorongoro Crater). Gestation 65–75 days. Litters average 2–3, exceptionally to 6. Kittens independent at 6–8 months. MORTALITY Known predators include Lion, Leopard, Nile Crocodile and domestic dogs. Predation by Martial Eagle on kittens recorded. LIFESPAN 11 years (s) in the wild, 20 in captivity. Status and Threats Conversion of wetland and grassland by draining, burning and overgrazing by livestock is a significant threat. Popular in the local fur trade in NE Africa and W African Sahel belt, and for fetish and traditional use in South Africa. Sport-hunted with few restrictions in Tanzania and southern Africa. May benefit from conversion of forest/woodland to grassland; expanding or recolonising range in Gabon, C Namibia, and C and NW South Africa. CITES Appendix II; Red List LC (global), CR (N Africa), population trend Stable.

CARACAL
Caracal caracal

Plate 7

HB 61–103cm, 62.1–108cm; T 18–34cm; W 6.2–15.9kg, 7.2–26kg

Uniformly coloured, pale sandy brown to brick red, unmarked except for faintly spotted undersides. Ears have conspicuous silvery-black backs and long black tufts. Melanistic cases are actually dark chocolate brown. Formerly classified as a lynx, but they are not closely related. Distribution and Habitat Africa (except for true desert and rainforest), the Middle East and SW Asia. Prefers dry woodland savannah, dry forest, grassland, coastal scrub, semi-desert and arid mountainous habitat. Sometimes inhabits evergreen and montane forests, exceptionally to 3,300m. Tolerates agricultural landscapes with cover. Feeding Ecology Formidable hunter, recorded killing adult Bushbuck, Springbok and Impalas, but most prey weighs <5kg; small rodents, hyraxes, hares, Springhare and birds are the most important. An estimated 2,920–3,285 rodents are eaten by each adult annually (Sariska TR, India). Reptiles comprise 12–17% of the diet (West Coast NP, South Africa); amphibians, fish and invertebrates are occasionally consumed. Readily kills small untended livestock, which comprises up to 55% of the diet in farming areas in southern Africa. Occasionally hoists kills into trees, and readily scavenges. Social and Spatial Behaviour Solitary. Adults maintain enduring home ranges with exclusive core areas and overlap at the edges. Female territories 3.9–26.7km2 (South Africa) to an average of 57km2 (Israel), compared with male territories of 5.1–65km2 (South Africa) to an average of 220km2 (Israel). An adult Kalahari male used 308km2, and an adult Saudi male used 865km2 (Harrat-al-Harrah PA). Density poorly known; 15/100km2 (protected area with abundant rodents and no large carnivores, Postberg NR, South Africa). Reproduction and Demography Breeds year-round, but births peak October–February (South Africa), and November–May (E Africa). Gestation 68–81 days. Litters average 2–3, exceptionally to 6. Kittens independent at 9–10 months. MORTALITY Most known mortality is anthropogenic; occasionally killed by larger carnivores, including domestic dogs in rural areas; Black-backed Jackals recorded killing kittens. Infanticide by males documented rarely. LIFESPAN 19 years in captivity. Status and Threats Habitat degradation, loss of prey and human hunting are significant threats in C, W and N Africa, and Asia, where Caracals are rare. An estimated 50–100 Caracals are trafficked annually from Somalia to the Gulf States for the pet trade. Persecuted intensely on livestock land in E and southern Africa, but resilient and difficult to extirpate there. Sport-hunted with few restrictions in E and southern Africa. CITES Appendix I – Asia, Appendix II – elsewhere; Red List LC, population trend Unknown.

AFRICAN GOLDEN CAT
Caracal aurata

Plate 7

HB 61.6–101cm; T 16.3–37cm; W 5.3–8.2kg, 8–16kg

Two distinct colour morphs, red-brown and grey, ranging from heavily spotted to plain in either. There is some intergradation between morphs, and melanism occurs. Does not change colour as is sometimes claimed. Distribution and Habitat Endemic to equatorial Africa. Strongly associated with undisturbed moist forests to 3,600m, including alpine bamboo forest, dense coastal forest and riverine forest strips in woodland savannah. Occurs in banana plantations inside forest, and in abandoned logged areas with secondary undergrowth. Avoids open and dry habitats. Feeding Ecology Eats a wide variety of small prey typically weighing 1.5–3.6kg; birds, shrews, rodents and small forest duikers are the most important. Actively hunts forest monkeys, but hunting is mainly terrestrial; scavenging of eagle kills on the forest floor may account for most primate occurrences in the diet. Reported to raid poultry in villages and to scavenge from wire snares. Social and Spatial Behaviour Unstudied and very poorly known. Solitary. Adults urine-mark and leave faeces exposed on trails, suggesting territorial behaviour. Reproduction and Demography Gestation 75 days. Litter size 1–2. Kittens weaned at around 6 weeks (captivity). Sexual maturity in captive animals 11 months (s) and 18 months (s). MORTALITY Poorly known; Leopard is a confirmed predator. LIFESPAN 12 years in captivity. Status and Threats Thought to be naturally rare. Forest-dependent; many W and E African moist forests are now heavily degraded and converted to savannah. Bushmeat hunting in W and C Africa heavily impacts prey species, and African Golden Cats are killed frequently in some areas for bushmeat and fetish markets. CITES Appendix II; Red List VU, population trend Decreasing.

Plate 7

MARGAY
Leopardus wiedii

Plate 8

TREE OCELOT

HB 47.7–62cm, 49–79.2cm; T 30–52cm; W 2.3–3.5kg, 2.3–4.9kg

Lightly built, resembling a small, lean Ocelot, but with a proportionally much longer tail, rounded head and distinctive large eyes. Melanism is very rare, with only two apparent records from camera-trap photos in Colombia (2013) and Costa Rica (2014). Distribution and Habitat From N Mexico throughout Central and South America to N Argentina, E Paraguay and N Uruguay. One specimen from S Texas around 1850 is the only US record, possibly a former captive given that the arid habitat is very atypical for the species. Forest-dependent, typically to 1,500m, exceptionally to 3,000m in the Andes; more closely associated with forest habitats than any other Neotropical cat. Avoids converted landscapes except for dense plantations, e.g. of coffee, cocoa, eucalyptus and pine. Feeding Ecology Most prey weighs <200g, and mainly comprises small rodents, shrews and mouse opossums. Larger prey includes Southern Opossum, cavies, agoutis, pacas and Brazilian Rabbits. Birds, including chachalacas and guans, herptiles, invertebrates and small amounts of fruit are eaten. Forages terrestrially and arboreally: a spectacularly acrobatic climber able to hunt the most agile prey, including small primates, although foraging is likely to be mainly ground-based. Occasionally kills domestic poultry. Mainly nocturnal, with peak activity 2100–0500. Social and Spatial Behaviour Poorly known, with very limited telemetry data. Solitary. Range sizes 0.9–20km2 (s) and 1.2–15.9km2 (s). Based on camera-trapping, reaches lower densities than Ocelot, although there are few estimates; 12 cats/100km2 estimated for protected montane pine–oak forest, C Mexico. Reproduction and Demography Unknown from the wild. In captivity, known for surprisingly low reproductive rates, with protracted gestation and small litters. Aseasonal (captivity). Gestation 76–84 days. Litter size usually 1, rarely 2. MORTALITY Poorly known. LIFESPAN 24 years in captivity. Status and Threats Strongly forest-dependent and responds poorly to forest conversion, its main threat. Formerly heavily hunted for fur (at least 125,547 skins were legally exported 1976–85), from which it may be slow to recover given its low reproductive potential. Localised illegal hunting and persecution for killing poultry is likely to have significant effects on populations in areas where there is existing pressure on habitat. CITES Appendix I; Red List NT, population trend Decreasing.

OCELOT
Leopardus pardalis

Plate 8

HB 69–90.9cm, 67.5–101.5cm; T 25.5–44.5cm; W 6.6–11.3kg, 7–18.6kg

Latin America’s third-largest cat. Powerfully built, with stocky limbs and a relatively short tail that rarely reaches the ground. Fur is creamy buff, tawny, cinnamon, red-brown or grey, with white underparts, and very richly marked with highly variable combinations of open and solid black blotches, streaks and rosettes with russet-brown centres. Simple solid spots or blotches usually cover the lower legs, and the tail has partial or complete black rings and a black tip. Melanism is unknown. Distribution and Habitat N Mexico to S Peru, N Argentina and SE Brazil, including on Trinidad and Isla de Margarita, Venezuela. Two relict populations occur in extreme SE Texas, numbering 50–80. Five individuals (4 males, 1 unidentified) documented in extreme S Arizona 2009–15; it is unknown if a resident breeding population occurs. Absent from Chile and now thought extinct in Uruguay. Inhabits a wide range of habitats but always with dense cover, from arid scrub to all kinds of dry and moist forest, usually below 3,000m. Tolerant of modified habitat provided there is dense vegetation and prey, e.g. agricultural landscapes with extensive brush, such as fallow cultivation or rice fields with forest edges. Avoids very open habitat but readily hunts in pasture and grasslands close to cover, especially at night. Feeding Ecology A flexible generalist with a wide diet, capable of killing large prey such as sloths, tamanduas, howler monkeys, Crab-eating Raccoons, coatis, and juvenile peccaries and deer, but mostly eating small rodents and mid-sized vertebrates, e.g. agoutis, pacas, squirrels, opossums, armadillos and iguanas. Other relatively common prey includes rabbits, tree porcupines, tamarins and squirrel monkeys, as well as birds such as large guans, macaws and seriemas. Readily consumes aquatic and semi-aquatic prey, including fish, amphibians and crustaceans, indicative of the species’ ability to inhabit inundated habitats. Incidental prey includes bats, lizards, snakes, small turtles, caimans (presumably hatchlings) and arthropods. Sometimes kills poultry, otherwise not considered dangerous to stock. Foraging is chiefly nocturno-crepuscular and terrestrial, although Ocelots are adept climbers that sometimes hunt in trees, e.g. a young Mantled Howler Monkey (Isla Barro Colorado, Panama). Scavenges, including from refuse piles left by people fishing, and sometimes caches large carcasses by covering with debris. Social and Spatial Behaviour Solitary and territorial. Male ranges (average 5.2–90.5km2) overlap multiple female ranges (average 1.3–75km2). Ranges are smallest in the Brazilian Pantanal and largest in cerrado savannah (Emas NP, Brazil). Ocelots appear to reach higher densities than all smaller sympatric felids, reaching very high numbers in good habitat; density estimates include 2.3−3.8/100km2 (tropical pine forest, Belize), 13–19/100km2 (Atlantic forest, Brazil), 26/100km2 (tropical rainforest, Belize) and 52/100km2 (dry Chaco–Chiquitano forest, Bolivia). Reproduction and Demography Unexpectedly low reproductive rates, with a fairly long gestation, very small litters and long inter-litter intervals; lifetime reproductive output per female is similar to much larger felids. Aseasonal. Gestation 79–82 days. Litter size 1–2, exceptionally 3 (only known in captivity). Kittens independent at 17–22 months. MORTALITY Annual mortality (Texas) is 8% (resident adults) to 47% (dispersers). Predators include large cats, Coyote and domestic dogs; confirmed, rare records by Boa Constrictor and American Alligator. LIFESPAN 20 years in captivity. Status and Threats Widespread and often abundant, but reliant on dense habitat and has a low reproductive potential. Vulnerable to habitat loss, illegal hunting and persecution for depredation; all are prevalent over much of the range. Texan Ocelots die mainly from anthropogenic factors, especially as roadkills. CITES Appendix I; Red List LC, population trend Decreasing.

Plate 8

NORTHERN TIGRINA
Leopardus tigrinus
SOUTHERN TIGRINA
Leopardus guttulus

Plate 9

ONCILLA, LITTLE-SPOTTED CAT, TIGER CAT

HB 43–51.4cm, 38–59.1cm; T 20.4–42cm; W 1.5–3.2kg, 1.8–3.5kg

Considered a single species until 2013, tigrinas are now thought to represent a complex of closely related ‘cryptic’ species, between which the phylogenetic and geographic borders are poorly understood. Currently regarded as 2 distinct but very similar species that diverged at least 100,000 years ago, based mainly on genetic data. Two additional species have been proposed but lack sufficient evidence to be widely accepted: L. pardinoides in NW South America; and L. oncilla, an isolated population in Central America. The 2 tigrina species show no genetic evidence of recent interbreeding. However, Southern Tigrinas hybridise with Geoffroy’s Cats in the wild; Northern Tigrinas do not hybridise with Geoffroy’s Cats yet they show evidence of past, historical hybridisation with the Colocolo. Northern and Southern tigrinas are extremely similar in appearance. Very small, slender, lightly built cats about the size of a young, lean domestic cat. Fur colour pale to dark buff or ochre, marked with orderly rows of black or dark brown dabs, blotches or small rosettes with a coffee-brown or reddish centre. Northern Tigrinas apparently tend towards lighter colouring with smaller rosettes than Southern Tigrinas, but wide variation occurs in both species, and distinct morphological differences have yet to be established. Melanism occurs in both species. Distribution and Habitat Northern Tigrina: N South America, from Venezuela and Colombia through E Peru and NE Bolivia to C Brazil, with a disjunct population in the Cordillera Central of Costa Rica and extreme W Panama. Southern Tigrina: SE and S Brazil, SE Bolivia, E Paraguay and probably extreme NE Argentina. The boundary between the two appears to be C Brazil. Both inhabit a broad range of habitats, including all types of forest, woodlands, wet and dry savannahs, arid scrublands and coastal restinga (scrub on sandy beaches), from sea-level to at least 3,200m. Northern Tigrina recorded at 3,626m (Costa Rica) to 4,800m (Colombia). In Central America, Northern Tigrinas are restricted to oak-dominated cloud and elfin forests above 1,000m. Tigrinas are apparently very rare or absent from Amazon Basin lowland rainforest. Both species tolerate habitats close to people, including rangelands, plantations, agricultural mosaics and peri-urban areas even near large cities, provided there is dense cover. Feeding Ecology Prey typically weighs <1kg and mainly comprises rodents, shrews, small opossums, birds, eggs, reptiles and invertebrates. Mid-sized diurnal lizards (ameivas and small iguanas) dominate the diet in semi-arid caatinga, NE Brazil, where rodent densities are low. Thought to forage mainly terrestrially and nocturno-crepuscularly depending on prey activity (and possibly the presence of Ocelots, which might depress their density), e.g. Northern Tigrinas are more diurnal in caatinga, reflecting reliance on diurnal reptiles. Tigrinas rarely take poultry. Social and Spatial Behaviour Solitary. Range size 0.9–25km2 (s) and 4.8–17.1km2 (s). Density estimates 0.01/100km2 (lowland Amazon forest) to 1–5/100km2. Reproduction and Demography Unknown from the wild. Gestation 62–76 days. Litter size typically 1, rarely 2. MORTALITY Poorly known. Single records each of deaths to Ocelot predation and heartworm disease. Domestic dogs are likely to be a significant predator in anthropogenic landscapes. LIFESPAN 17 years in captivity. Status and Threats Both species appear to be naturally rare, elevating their vulnerability to threats, but their status is poorly known. They have become locally extinct where dense submontane forest has been converted for agriculture (Costa Rica and the Andes). Formerly heavily hunted for fur trade; some hunting for local fur demand still occurs. Localised killing for furs, by dogs, in retaliation for killing poultry and on roads probably impacts populations close to people. Both species: CITES Appendix I; Red List VU, population trend Decreasing.

GEOFFROY’S CAT
Leopardus geoffroyi

Plate 9

HB 43–74cm, 44–88cm; T 23–40cm; W 2.6–4.9kg, 3.2–7.8kg

Largest of the South American temperate zone small felids, reaching the size of a large domestic cat. Yellow-brown to silver-grey with small, solid dab-like spots, becoming elongated blotches on nape, chest and lower limbs; southern, temperate animals are typically paler. Melanism is common in Uruguay, SE Brazil and E Argentina, but rare elsewhere. Hydridises with Southern Tigrina in S Brazil. Distribution and Habitat C Bolivia to Uruguay and S Brazil, and most of Argentina. Inhabits subtropical and temperate brushland, forest, semi-arid scrub, pampas grassland and marshland, from sea-level to 3,300m in the Andes. Absent from tropical and temperate rainforest. Tolerant of disturbed habitats, including ranchland and conifer plantations. Feeding Ecology Eats small rodents such as grass mice, rice rats and cavies, as well as small birds and herptiles. Occasionally takes Six-banded Armadillo, tree porcupines and small opossums. Introduced European Hare dominates the diet in Patagonia. Large waterbirds, including cormorants, ibis, coots, ducks, Chilean Flamingo and Coscoroba Swan are primary prey in coastal lagoon habitat (Mar Chiquita, Argentina). Mostly nocturno-crepsucular and terrestrial, but swims readily, taking Coypu, marsh rats, frogs and fish at the water’s edge. Occasionally recorded caching large kills, e.g. hares, in trees and burrows. Raids domestic poultry; sheep are probably scavenged. Social and Spatial Behaviour Solitary. Deposits faeces in trees, creating arboreal middens. Range sizes average 1.5–5.1km2 (s) and 2.2–9.2km2 (s). Density estimates 4/100km2 (Argentine pampas during a prey shortage) to 45–58/100km2 (Argentine ranchlands); 139/100km2 estimated for protected Argentine scrubland may be an overestimate. Reproduction and Demography Seasonal in temperate southern range (possibly elsewhere), with most births December–May. Gestation 62–78 days. Litter size 1–3. MORTALITY Starvation and high parasite loads elevate mortality during droughts. Puma is a known predator. LIFESPAN 14 years in captivity. Status and Threats Widespread and abundant in good habitat, including some anthropogenic landscapes, but the species has disappeared where habitat is converted to agricultural monocultures. Often killed by vehicles, domestic dogs and for attacking poultry. CITES Appendix I; Red List LC, population trend Stable.

Plate 9

GUIÑA
Leopardus guigna

Plate 10

GUIGNA, KODKOD

HB 37.4–51cm, 41.8–49cm; T 19.5–25cm; W 1.3–2.1kg, 1.7–3kg

Tiny cat, grey-brown to russet-brown with small, dark dab-like spots coalescing into irregular lines on the back and nape. Face distinctively marked with dark stripes under the eyes bordering the muzzle, resembling that of a Puma kitten. Melanism is common, sometimes with rich brown (rather than black) extremities on which the markings are obvious. Distribution and Habitat C and S Chile, including Chiloé Island, and marginally in adjacent Argentina. Strongly associated with dense temperate habitats, especially evergreen forest, montane forest, thicket and scrubland. Avoids open land and plantations with little understorey, but uses secondary forest and exotic plantations, as well as forested ravines and coastal forest strips in cleared habitat. Feeding Ecology Hunts small rodents and marsupials, e.g. Monito del Monte, ground-foraging birds such as tapaculos, ovenbirds, thrushes and lapwings, and small reptiles and insects. Fishes for cavity-nesting bird chicks, e.g. rayaditos and wrens, in artificial nestboxes (and very likely natural cavities). Regularly kills free-range domestic poultry in fragmented human-dominated landscapes, e.g. Chiloé Island, and is considered a pest. Reports of goat killing and hunting in groups are implausible. Cathemeral. Social and Spatial Behaviour Solitary. Ranges overlap considerably, including in core areas, suggesting limited territoriality. Range size 0.6–2.5km2 (s) and 1.6–4.4km2 (s). Density estimates from radio-telemetry, counting adults and subadults, 1–3.3/km2. Reproduction and Demography Unknown from the wild; cold winters possibly drive seasonal breeding. Gestation 72–78 days. Litter size 1–3. MORTALITY Poorly known; where studied, humans are the main cause of death. LIFESPAN 11 years in captivity. Status and Threats Very restricted distribution and dependent on dense habitat. Forest loss from agriculture and pine plantations has reduced its range to many small, fragmented populations, which are further threatened by illegal killing, mainly over poultry depredation. CITES Appendix II; Red List VU, population trend Decreasing.

COLOCOLO
Leopardus colocolo

Plate 10

PAMPAS CAT

HB 42.3–79cm; T 23–33cm; W 1.7–3.7kg

Highly variable, ranging from smoky grey to dark rusty brown, with little or no spotting to rich russet-coloured blotches on the body. Based on morphology, the species clusters into 3 major groups: ‘Colocolo’ in Chile, W side of the Andes; ‘Pampas Cat’ in Colombia to S Chile, E side of the Andes; and ‘Pantanal Cat’ in Brazil, Paraguay and Uruguay. They are mistakenly sometimes considered separate species; all forms intergrade, and genetic data indicate only moderate differences. Melanism is recorded from Brazil and Peru. Distribution and Habitat S Colombia to S Chile, extending into C Brazil, Paraguay and Uruguay. Occupies more habitat types than any other Latin American felid, including pampas and cerrado grassland, woodland savannah, marshland, open forest, cloud forest, semi-arid desert and Andean steppes to 5,000m. Does not occur in rainforest. Tolerates plantations and agricultural habitat with cover, e.g. maize cropland. Feeding Ecology Focuses mainly on small mammals, especially tuco-tucos, mountain viscachas, chinchilla rats, leaf-eared mice, rats and introduced European Hare. Other notable prey includes flamingos (probably chicks), tinamous, Magellanic Penguin chicks, small reptiles, eggs and beetles. Kills domestic poultry. Foraging is generally terrestrial and nocturnal, but varies with the region; almost entirely diurnal in the Brazilian cerrado, perhaps due to the presence of nocturnal large cats. Scavenges from carcasses, including those of livestock, Vicuña and Guanaco. Social and Spatial Behaviour Solitary. Range size 3.7–37km2, averaging 19km2 (Emas NP, Brazilian cerrado) and 4.7–55.3km2 (Argentina–Bolivia border region). Density estimates 11.3–17.6/100km2, (Argentine espinal forest-grasslands) and 20–78/100km2 (3 surveys in Argentine high Andes, upper figures are possibly an overestimate). Reproduction and Demography Unknown from the wild. Gestation 80–85 days. Litter size 1–3. MORTALITY Poorly known. Apparently frequently killed by domestic dogs in some locations, e.g. NW Argentina. LIFESPAN 16.5 years in captivity. Status and Threats Widely distributed, with a broad habitat tolerance, and often fairly rare in camera-trap surveys compared to sympatric felids. Killed for raiding poultry and vulnerable to shepherds’ dogs, especially in open Andean habitats. In rural areas, killed for religious ceremonial uses in which the skin or a stuffed cat is believed to confer fertility and productivity on domestic livestock and crops. CITES Appendix II; Red List NT, population trend Decreasing.

ANDEAN CAT
Leopardus jacobita

Plate 10

ANDEAN MOUNTAIN CAT

HB 58–75cm; T 38–46cm; W 4.6–5.8kg

Silver-grey marked with russet blotches on the body that darken to rich grey-brown on the face, limbs and tail. Tail very thick and bushy, with distinctive thick banding that becomes paired brown or russet rings, often with mid-brown centres, towards the tip. Distribution and Habitat S Peru to NW Argentina; restricted to high Andean habitats mostly at 3,000–5,100m. Has been recorded from the Patagonian steppe at 1,800m (Mendoza, Argentina) and 600–700m (Neuquén, Argentina). Occurs only in semi-arid to arid treeless habitats with rocky slopes and cliffs, and associated shrubland and grassland. Feeding Ecology Specialises in high-altitude rodents, especially 2 species of mountain viscacha, as well as mice, leaf-eared mice and chinchilla rats; cavies, European Hare and tinamous are also recorded prey. Mostly nocturnal, with crepuscular activity peaks in high-altitude areas reflecting activity patterns of viscachas. Sometimes scavenges from the carcasses of dead ungulates. Social and Spatial Behaviour Solitary. Range size 20–65km2 (Argentina–Bolivia border region), and 47.1km2 for a female in the Bolivian altiplano monitored for 4 months. Density estimates 7–12/100km2 (Argentine high Andes). Reproduction and Demography Poorly known. Kittens have been observed October–April, suggesting seasonal breeding with spring/summer births. Litter size 1–2. MORTALITY and LIFESPAN Poorly known. People and dogs are the primary known sources of mortality. Status and Threats Status virtually unknown. During surveys, evidence of Andean Cat is found far less frequently than that of Colocolo and other carnivores, suggesting the species is naturally rare. It has a very restricted distribution and narrow habitat preference, which is vulnerable to livestock grazing and agriculture. This may also impact on prey numbers, especially combined with hunting of prey, particularly viscachas, which is considered a serious threat. Killed for religious beliefs (see Colocolo) and for suspected poultry and livestock killing; sometimes killed by herders and their dogs, e.g. Neuquén Province, Argentina, even though it does not prey on stock. CITES Appendix I; Red List EN, population trend Decreasing.

Plate 10

EURASIAN LYNX
Lynx lynx

Plate 11

HB 85–130cm, 76–148cm; T 12–24cm; W 13–21kg, 11.7–29kg

Largest lynx. Colour varies from silver-grey and tawny to red-brown, with highly variable spotting, including unspotted, small coin-like spots, large dab-like spots and elongated brown rosettes. Coloration varies considerably between and within populations. Distribution and Habitat Fenno-Scandinavia, Russia (75% of range), China and temperate C Asia, with scattered populations in E and W Europe. Inhabits mainly forest, montane areas with cover, cold semi-desert, tundra, open woodland and scrub. Occurs to 4,700m in the Himalayas, exceptionally to 5,500m. Feeding Ecology In contrast to other lynx, hunts mainly small to medium-sized ungulates. The most important prey across much of its range is European and Siberian roe deer, followed by Chamois, musk deer and juveniles of Red Deer, Sika Deer, Moose, ibex and Wild Boar. Ungulates are especially important prey in winter, when snow elevates vulnerability to predation: exceptional kills of adult Red Deer occur in deep snow. Hares, small rodents, squirrels, marmots and birds increase in importance during spring and summer. In far northern forests, where ungulates are less common, Mountain Hares and European Hares are the most important prey year-round. Kills livestock and poultry, including frequent predation on semi-domestic Reindeer in Finland, Norway and Sweden. Mainly nocturno-crepuscular, but may be cathemeral, especially in winter and during the breeding season. Scavenges, and sometimes caches large carcasses with a covering of ground debris. Social and Spatial Behaviour Solitary. Male ranges are larger than female ranges, and overlap more extensively. Both sexes demarcate territorial boundaries with urine, scent and faecal marks, but ranges are generally too large to permit exclusivity, except among females with small kittens. Range sizes increase from south to north, reflecting prey availability; 98–1,850km2 (s) and 180–3,000km2 (s). Average range size in W and C Europe 106–264km2 (both sexes combined), compared with 307–1,515km2 in Scandinavia. Largest recorded ranges are in Norway. Density estimates include 0.25/100km2 (S Norway) to 1.9–3.2/100km2 (Poland). Reproduction and Demography Seasonal. Mates February–mid-April; births May–June. Oestrus 3–5 days; gestation 67–74 days. Litter size 1–4, very rarely 5, typically 2. Independent at 9–11 months, with dispersal occurring before the mother’s next litter. Females first breed at 22–24 months, males at 3 years. MORTALITY Kitten mortality usually at least 50%; 59–60% of Swiss kittens die before independence. Natural adult mortality low, only 2% annually in Scandinavia, but anthropogenic factors increase that by a factor of eight; 44–60% of subadults die during dispersal (Switzerland). Predation occurs occasionally by Grey Wolf, Tiger and Wolverine (young animals). LIFESPAN 18 () and 20 () years in the wild, 25 in captivity. Status and Threats Considered secure, with large areas of its massive range still intact, especially in Russia (where the population is estimated at 30,000–35,000), Mongolia and China. Extirpated from most of W and C Europe, where remaining populations are small and isolated, but recovering and expanding in W Europe, e.g. the Alps. No longer legally hunted for fur except in Russia (~1,000 lynx/year), although illegal trade is widespread in its Russo-Asian range. Sport hunting is legal in much of the range; the highest quotas are in Russia, Estonia, Finland, Latvia, Norway and Sweden. CITES Appendix II; Red List LC, population trend Stable.

IBERIAN LYNX
Lynx pardinus

Plate 11

SPANISH LYNX, PARDEL LYNX

HB 68.2–75.4cm, 68.2–82cm; T 12.5–16cm; W 8.7–10kg, 7–15.9kg

Tawny grey to reddish brown, with solid spotting or blotches that sometimes break up into freckling. Both sexes have a prominent white facial mane with black streaks, and the ears have long black tufts. Tail ends in a black tip. Distribution and Habitat Restricted to 3 populations on the southern Iberian Peninsula: the largest, a meta-population comprising a main population with 2 satellite reintroduced populations, Sierra Morena mountains, S Spain; a population in Doñana NP (S Spain); and a small, reintroduced population in Vale do Guadiana NP, SE Portugal. Occurs in dense mosaics of forest, thicket, brushland and Mediterranean scrub, with open pastures and edges favoured for hunting. Shuns agricultural land and exotic plantations, but uses pine plantations for dispersal. Feeding Ecology Highly reliant on European Rabbit, which comprises 75–93% of the diet, depending on the location and season. Cannot live in areas without abundant rabbits, and requires an estimated 277/year for a female without kittens, up to 379/year for a male. Incidental prey includes small rodents, hares and birds, including ducks, geese, partridges, magpies and pigeons. Juvenile Red Deer and Fallow Deer are sometimes killed during autumn and winter. Kills other carnivores, including Red Fox, Egyptian Mongoose, Common Genet and feral domestic cats, but these are rarely eaten and probably mainly killed as competitors for rabbit prey. Kills rabbits by biting the skull; larger prey such as deer are killed by suffocation. Does not kill livestock or poultry, due largely to very limited opportunity in its remaining range. Covers larger kills with leaf litter and debris to consume over a number of days. Social and Spatial Behaviour Solitary and territorial. Each male range overlaps 1 and sometimes 2 female ranges, with intra-sexual overlap at the edges and exclusive core areas. Territorial fights are occasionally fatal. Range size 8.5–24.6km2, averaging 12.6km2 (s), and 8.5–25km2, averaging 16.9km2 (s). Density estimates 10–20/100km2 with moderate rabbit densities to 72–88/100km2 in highly protected enclaves with extremely abundant rabbits. Reproduction and Demography Seasonal. Mating usually January–February, with births peaking March and occasional births April–June. Litter size 2–4, averaging 3. Kittens independent at 7–8 months, remaining in their natal range until dispersal at 18–20 months. Females can breed at 2 years, but usually first give birth in the wild at 3 years, and breed until age 9. MORTALITY Kitten mortality around 33%; 2 kittens usually survive from most litters of 3. Adult mortality around 10% for resident animals insulated from human factors, but anthropogenic mortality is now the major cause of Iberian Lynx deaths. LIFESPAN 10 years in the wild, 14 in captivity. Status and Threats The world’s most endangered felid in terms of total population, which numbers approximately 400 adults in 2 isolated populations in Spain, plus 10 adults reintroduced to SE Portugal (2014–15; kittens documented 2016). Decline driven by extensive habitat conversion of forest to exotic plantations, combined with disease epidemics in rabbits and direct human killing of lynxes. A massive conservation effort has produced recovery from fewer than 150 adults in 2002, and they were downlisted from Critically Endangered in 2015. Anthropogenic killing has declined but remains a serious threat, responsible for 75% of lynx deaths, mostly by illegal trapping, shooting and roadkills. CITES Appendix I; Red List EN, population trend Increasing.

Plate 11

BOBCAT
Lynx rufus

Plate 12

BAY LYNX, RED LYNX

HB 50.8–95.2cm, 60.3–105cm; T 9–19.8cm; W 3.6–15.7kg, 4.5–18.3kg

Various shades of grey to rusty brown, with markings ranging from very minimal spotting to large, Ocelot-like blotches. Bobtail has 3–6 dark half-stripes and a vivid white underside and tip, distinguishing the species from Canada Lynx. Melanism occurs occasionally, recorded mostly from SE USA. Distribution and Habitat Southern Canada, USA and N Mexico. Very wide habitat tolerance, including all forest types, brushland, scrub, prairies, semi-desert and mountainous terrain. Tolerant of farmland, agricultural land and peri-urban landscapes provided there is cover. Feeding Ecology Recorded killing adult deer weighing up to 68kg, but typical prey is hare-sized or smaller. Lagomorphs are key prey throughout its range, especially Snowshoe Hare, Eastern Cottontail, jackrabbits and Marsh Rabbit. White-tailed Deer, Mule Deer and Pronghorn are taken primarily as fawns, but northern Bobcats kill more adults, especially in winter. Other prey includes rodents to the size of porcupines, smaller carnivores, opossums, birds, herptiles, fish, arthropods and eggs. Kills sheep, goats and poultry, although problems are usually localised. Rarely kills small pets in peri-urban areas (small dogs and domestic cats are not recorded in the diet). Forages mostly on the ground. Generally nocturnal with crepuscular activity peaks. Sometimes caches carcasses with a covering of dirt or snow to consume over time, e.g. up to 14 days for an adult deer kill. Eats carrion: road-and winter-killed deer are important food sources, especially in northern winters. Social and Spatial Behaviour Solitary and territorial. Range size and overlap decrease with increasing prey availability. Territorial fights are occasionally fatal. Average range size 1–2km2 (Alabama; California; Louisiana; Oregon) to 86km2 (Adirondacks, New York) for females, and 2–11km2 (Alabama; California; Louisiana; Oregon) to 325km2 (Adirondacks) for males. Ranges contract during prey peaks, especially of hares and rabbits. Nightly movements are as large as 20km. Density estimates include 4–6/100km2 (Minnesota), 20–28/100km2 (Arizona, Nevada), exceptionally to >100/100km2 (e.g. coastal California, when protected from hunting). Reproduction and Demography Weakly seasonal. Births occur year-round, but peak spring–summer, strongly so in northern areas. Gestation 62–70 days. Litter size averages 2–3, exceptionally to 6. Weaning at 2–3 months. Kittens independent at 8–10 months. Females can breed at 9–12 months, but usually first give birth after 24 months. MORTALITY Kitten mortality fluctuates extensively, depending mainly on prey numbers, e.g. 29–82% mortality in Wyoming in different years. Adult mortality 20–33% for unharvested populations to 33–81% for hunted populations. Mostly killed by humans, as well as by winter starvation, predation by Puma, Coyote and domestic dogs, and episodic disease outbreaks in dense populations. LIFESPAN 16 years in the wild, 32.2 in captivity. Status and Threats Widespread, resilient to human pressures and secure in most of its range, with >1 million estimated for the US alone. None the less, some populations are exposed to intense hunting pressure and Bobcats are vulnerable to overharvesting. Around 40,000–50,000 are legally killed in the USA and Canada annually, mainly for skins; globally, it is the most heavily traded felid species for fur. Persecuted for supposed livestock depredation, e.g. Mexico, and 2,000–2,500 are killed annually in legal control in the USA. CITES Appendix II; Red List LC, population trend Stable.

CANADA LYNX
Lynx canadensis

Plate 12

CANADIAN LYNX

HB 76.2–96.5cm, 73.7–107cm; T 5–12.7cm; W 5–11.8kg, 6.3–17.3kg

Uniformly coloured, typically buff-grey with silver or bluish frosting in winter, and brownish in summer, unspotted or lightly spotted on the limbs. Tail shorter than Bobcat’s, with a completely black tip. Lynx–Bobcat hybrids occasionally occur where the 2 species overlap in Maine, Minnesota and New Brunswick. Distribution and Habitat Most of Canada south of the treeline and some US border states, south to Utah. Reintroduced successfully to S Colorado and unsuccessfully to New York. Closely tied to dense boreal and coniferous forests; rarely uses open habitat. Feeding Ecology Strongly dependent on Snowshoe Hare, which comprises 35–97% of the diet locally and seasonally. Northern hare populations cycle every 8–11 years, sometimes spectacularly, from 2,300/km2 to 12/km2. Lynx numbers are closely linked, lagging 1–2 years behind. Lynx switch prey during declines and during summer. Southern populations (which experience weak or non-existent hare cycles) have more diverse diets year-round, although Snowshoe Hare remains the primary prey. Other important prey includes American Red Squirrel, other rodents, small birds, and game birds like grouse and ptarmigans. Ungulate lambs, especially of Caribou, are sometimes killed, but most ungulates are scavenged. Rarely kills livestock or poultry. Hunting is mainly crepuscular, nocturnal and chiefly terrestrial. Occasionally caches prey by covering it with snow or leaves. Social and Spatial Behaviour Solitary and probably territorial, but spatial behaviour varies extensively depending on Snowshoe Hare availability. Southern populations with stable but low densities of hares tend to maintain large, enduring home ranges with high overlap between neighbours. Northern populations maintain smaller and possibly more exclusive ranges during hare peaks, but ranges expand during declines, sometimes leading to nomadism. Average range size estimates include: 39–133km2 (s) and 69–277km2 (s) in southern populations; 13–18km2 (s) and 14–44km2 (s) in northern populations with high hare numbers; and up to 63–506km2 (s) and 44–266km2 (s) in northern populations with few hares. Density estimates include 2–4/100km2 during low hare density to 10–45/100km2 during peaks. Reproduction and Demography Seasonal. Mating March–May; births May–early July. Gestation 63–70 days. Litter size 1–8, with more females breeding and larger litters produced during hare peaks. Independent at 10–17 months. Females breed as early as 10 months during high hare years, but usually first breed at 22–23 months. MORTALITY Kitten survival is linked closely to hare numbers, reaching 60–95% mortality in poor years. Estimates of adult mortality include: 11–27% and up to >60% during hare shortages for unharvested populations; and 45–95% for harvested populations. Starvation and trapping by humans are responsible for most deaths. LIFESPAN 16 years in the wild, 26.9 in captivity. Status and Threats Generally widespread and secure, especially in Canada. Extirpated from most of its US distribution, and its range appears to be retreating north with forestry and climatic warming; suitable habitat has receded ~175km northwards in C Canada since the 1970s. On average, at least 11,000 Canada Lynx are legally harvested annually, most in Canada and Alaska; they are vulnerable to overharvesting during Snowshoe Hare declines. CITES Appendix II; Red List LC, population trend Stable.

Plate 12

JAGUARUNDI
Herpailurus yagouaroundi

Plate 13

EYRA

HB 53–73.5cm, 48.8–83.2cm; T 27.5–59cm; W 3.5–7kg, 3–7.6kg

Uniformly coloured with 2 distinct, highly variable morphs: red-brown (often with a bright white muzzle and chin), varying from light tawny to brick red; and grey, varying from pale slate grey to deep black-grey (often with a paler neck and head). Litters can include kittens of both colours. Body and head elongated and lean, with a long, slender tail, giving the impression of a mustelid in the field. Distribution and Habitat C Argentina to N Mexico, and formerly in the Rio Grande Valley, Texas, USA, but not confirmed there since 1986. Occurs from sea-level to 2,000m (occasionally to 3,200m) in all types of forest, scrub, chaparral, brush, dense grassland and pasture. Does well in human-modified or recovering habitat with cover and high rodent densities, e.g. pasture grassland, old fields and secondary forest. Feeding Ecology Makes kills to the size of an armadillo, but most prey weighs less than 1kg. Principal prey includes rats, mice and birds, especially ground-dwellers like tinamous and quail. Also eats small primates (rarely), cavies, rabbits, opossums, reptiles, fish and arthropods. Brocket deer are recorded in scats, but probably from scavenging. Occasionally raids poultry coops. Appears to be primarily crepuscular/diurnal; a capable climber, but most hunting takes place on the ground. Social and Spatial Behaviour Solitary. Exhibits typical felid marking behaviour and is presumably territorial, although radio-tracked individuals overlap extensively. Based on relatively few studies, range size is similar for females and males, or slightly larger for the latter: 12.1km2 (s) and 16.2km2 (s; Tamaulipas, Mexico); 1.4–18km2 (s) and 8.5–25.3km2 (s; Brazil). Ranges for 3 monitored Belizean cats were 20km2 (1 ) and 88–100km2 (2 s), the latter an unusually high figure probably indicating dispersers. Reproduction and Demography Poorly known; possibly weakly seasonal, although reports are contradictory. In captivity, gestation 72–75 days, and litters number 1–4 (average 1.8–2.3). Weaning begins at around 5–6 weeks. Sexual maturity at 17–26 months. MORTALITY Poorly known. Recorded predators include Puma, and domestic dogs near villages. LIFESPAN 10.5 years in captivity. Status and Threats Tolerant of human activity and seldom hunted for its unicolour fur. Persecuted for killing poultry, and common roadkill in some areas, but most populations are secure. Widespread and relatively common in South America. Endangered in Central America; considered Critically Endangered in the USA, but likely extinct there. CITES Appendix I – Central and North America, Appendix II – elsewhere; Red List LC, population trend Decreasing.

PUMA
Puma concolor

Plate 13

COUGAR, MOUNTAIN LION, PANTHER (FLORIDA)

HB 95–141cm, 107–168cm; T 57–92cm; W 22.7–57kg, 39–80kg (exceptionally to 125kg)

Uniformly coloured, ranging from light grey through tawny brown to brick red, with creamy-white underparts. Temperate Pumas tend to be larger, with paler, greyish coloration, while tropical individuals are smaller with richer reddish tones. Tail tip and backs of the ears are dark brown to black, and the white muzzle is bordered by black. Long tubular tail is distinctive in the field. Cubs have rich, dark brown spots that usually fade within the first year. Distribution and Habitat Relatively widely distributed in SW Canada, W USA and South America. More restricted in Mexico and Central America, and extirpated from E USA except for 100–120 in S Florida (‘Florida Panther’). Presence in El Salvador is uncertain, with no recent records. Very wide habitat tolerance provided there is vegetation or rocky terrain, including temperate and tropical forests, woodland, coastal and desert scrublands, and rocky desert, from sea-level to >4,000m (exceptionally to 5,800m in the Andes, S Peru). Mostly shuns open areas, but readily passes though marginal habitat. Lives close to humans provided cover and prey are available. Feeding Ecology Very broad diet encompassing arthropods to adult male Elk. Large kills are more common in temperate populations, where deer, Moose, Elk and wild sheep form the principal prey; Guanaco is a key prey species in Chile. Tropical Pumas focus more on smaller prey such as brocket deer, peccaries, Capybara, paca, agoutis and armadillos. Locally, the diet may be dominated by feral livestock, e.g. pigs (Florida) and wild horses (Nevada). Kills reptiles to the size of adult caimans and alligators, and birds such as Wild Turkey, various geese and rhea. Readily kills livestock, and sometimes takes domestic pets in peri-urban areas. Humans are rarely killed, 26 recorded fatalities (including 2 from rabies infection) in North America in 1890–2018. Hunting is mainly nocturno-crepuscular. Typically discards entrails of large kills before covering them in dirt or leaf litter, consuming the carcasses over a period of 3 days to 4 weeks (winter). Scavenges, although this usually represents a small amount of its intake. Social and Spatial Behaviour Broadly solitary and territorial, but with considerable overlap in ranges, and greater sociality in some populations, e.g. Greater Yellowstone Ecosystem resident adults have frequent non-agonistic interactions among non-related animals, including prolonged associations and sharing large carcasses for up to 5 days. Territorial fights are sometimes fatal, particularly in disturbed populations (such as those heavily hunted). Home range is 25–1,500km2, averaging 33km2 (Venezuela) to 685km2 (Utah) for females, and 60km2 (Venezuela) to 826km2 (Utah) for males. Pumas are superb dispersers: 6–32km (s) and 24–208km (s; Florida), to 12–99km (s) and 12–1,067km (s; Black Hills, South Dakota). In 2011, a South Dakota male was killed by a vehicle in Connecticut ~2,800km away. Density estimates: 0.3/100km2 (e.g. Utah; Texas) to 1–3/100km2 (Alberta; California; Utah; Wyoming), 3–4.4/100km2 (Pantanal, Brazil), 2.4–4.9/100km2 (rainforest, Belize), exceptionally to 7/100km2 (Vancouver Island). Reproduction and Demography Weakly seasonal. Births occur year-round, but typically peak in summer (Yellowstone NP; Canada) or spring (Florida). Oestrus lasts 1–16 days; gestation averages 92 days (range 82–98 days). Litter size averages 2–3 kittens, exceptionally to 6. Weaning at around 4–5 months. Cubs independent at around 18 months (range 10–24 months). Females first give birth at 18 months (typically >24 months) and males first breed at around 3 years. MORTALITY 36–58% of cubs die in their first year. Estimates of adult mortality include 9% (s) to 18% (s) in New Mexico, and 3% (s) to 61% (s) in NW Montana. Mostly killed by humans (legal hunting, roadkills, illegal killing); principal natural causes of death for adults are starvation, disease and hunting accidents. LIFESPAN 16 years in the wild, 20 in captivity. Status and Threats Tolerant of human activity, but extirpated from around 40% of its Latin American range and most of its eastern North American range. Habitat loss, combined with intense persecution in livestock areas, are the major threats, especially in Latin America. Legal sport hunting kills 2,500–3,500 Pumas per year in the USA, triggering population declines in some states. Road accidents are a major threat in Florida, e.g. 63% of 381 known deaths in 2000–17. Argentina is the only range state still paying bounties in Pumas, producing extremely excessive harvests of ~2,000 annually. CITES Appendix I – Nicaragua through Panama, Appendix II – elsewhere; legally hunted in Argentina, Canada, Mexico, Peru and USA (all states except California and Florida); Red List LC, population trend Decreasing.

Plate 13

CHEETAH
Acinonyx jubatus

Plate 14

HB 105–140cm, 108–152cm; T 60–89cm; SH 67–89cm, 74−94cm; W 21–51kg, 29–64kg

Yellow-blond fading to white underparts, with black coin-like spots and unique facial tear streaks. Saharan Cheetahs have very short, pale fur, ranging from brown-spotted beige to near white with faint cinnamon spots. So-called King Cheetahs are a recessive colour morph and may be born to normally spotted parents. Cubs have a fluffy smoky-grey mantle that dwindles by 4–5 months to a short mane on the shoulders, which is inconspicuous in most adults but often obvious in Asiatic Cheetahs. Mantle’s function is unclear, but it probably assists with camouflage and thermoregulation; mimicry of Honey Badger to deter predators, as is often claimed, is doubtful. Claws dog-like and lack fleshy claw-sheaths present in other cats, but are partially protractile. Claws appear in tracks, except for the sharp, strongly curved dewclaw, which is used for prey capture.

Distribution and Habitat

Relatively widely distributed in southern and E Africa, rare in W Africa, and extinct in N Africa except S Algeria and possibly W and NE Libya. Extinct in Asia except for ~50 in Iran. Favours woodland–savannah mosaics and open grassland, becoming sparser in dense humid woodland, e.g. Zambian miombo, and absent from rainforest. Tolerates arid habitats, including the deserts of C Iran (which experience winter snowfall), and the Namib and Sahara, but transient in the driest areas. Recorded to 3,500m (Mt Kenya, Kenya).

Feeding Ecology

Usually hunts ungulates weighing 10–60kg, particularly gazelles, Impala, Steenbok and duikers, occasionally specialising in prey up to size of Nyala (62–108kg); also juveniles of larger ungulates, including oryx, wildebeest, zebra and (rarely) buffalo and giraffe. Male coalitions take large prey, e.g. adult hartebeest, oryx and wildebeest. Asiatic Cheetahs eat mainly Urial and Persian Ibex following the widespread extirpation of gazelles. Hares are important prey in certain habitats, e.g. S Kalahari, and especially to recently independent young adults and in prey-depleted landscapes, e.g. Iran. Kills small untended livestock, but is easily deterred by people or dogs, and is not recorded killing humans. Hunting mostly diurnal to maximise visibility and avoid nocturnal competitors. Following a short and careful stalk, pursues prey for up to 500m at a maximum recorded speed of 105km/h. Prey is bowled over or pulled off balance using the dewclaw, and usually killed by suffocation. Around 30% (Serengeti NP, Tanzania) to 43.4% (S Kalahari, South Africa) of chases are successful, more so for small prey: 73–93% for hares (Kalahari and Serengeti NP), 86–100% for juvenile gazelles (Serengeti NP). Rarely defends kills, losing up to 13% (Serengeti NP), chiefly to Spotted Hyaena and Lion. Very rarely scavenges and seldom returns to carcasses previously fed upon.

Social and Spatial Behaviour

Unusual social system in which females are solitary and non-territorial, while male sociality and territoriality vary widely. Females occupy large home ranges that are not defended. Where prey is scarce or migratory, female ranges reach 1,500km2 (average 833km2; Serengeti NP) to 6,353.7km2 (average 1,400–1,836km2; Namibia). Areas with resident or abundant prey produce smaller ranges (e.g. 185–246km2; Kruger NP, South Africa), although females are non-territorial regardless. Dispersing females usually settle near their natal range, so that neighbouring females are often related. Males typically disperse further to avoid breeding with female relatives, and live in permanent coalitions of 2–4, usually littermates; around a third of Serengeti coalitions include an unrelated member. Coalitions defend territories where profitable, repelling rival males in fights that may be fatal. Serengeti coalitions target small areas with high female overlap averaging 37km2. Males in woodland habitats with resident females establish medium-sized territories, e.g. Kruger NP, 126km2 (3 coalition) to 195km2 (single ). Coalitions defend territories more successfully than loners, and single males are often nomadic ‘floaters’ with much larger home ranges, averaging 777km2 (Serengeti NP) to 1,829km2 (Namibia). A pair of Iranian males (likely nomadic) used 1,737km2 in 5 months of radio-tracking. Coalitions sometimes also float on large ranges (average 1,608.4km2; Namibia) as an alternative to territorial defence. Cheetahs naturally occur at low densities: 0.16/100km2 (Iran), 0.25–2/100km2 (Namibian farmland) and 0.5–2.30/100km2 (Kruger NP), to 2.5/100km2 (Serengeti NP). Seasonal densities exceptionally reach 20/100km2 when Cheetahs congregate temporarily in localised areas (Serengeti NP).

Reproduction and Demography

Cheetahs have a reputation for poor reproduction arising from the difficulties of breeding them in captivity, but wild Cheetahs are prolific. Aseasonal, although Asiatic Cheetahs apparently mate in winter (January–February) and give birth in spring (April–May). Gestation 90–98 days. Litter size typically 3–6, rarely to 8; litter of 9 from Kenya may have included an adoption. Inter-litter interval averages 20.1 months (Serengeti NP). Weaning begins at 6–8 weeks and is complete by 4–5 months. Cubs independent at 12–20 months (average 17–18 months) and disperse as a sibling group; females leave the group before sexual maturity, while males remain together. Females can conceive at 21–24 months; first give birth around 29 months (Serengeti NP) and can reproduce to 12 years. Males are sexually mature at 12 months, although they rarely breed before 3 years. MORTALITY 71% (S Kalahari) to 95% (Serengeti short grass plains) of cubs die before independence, most killed by other carnivores. Losses in the denning period are the highest. In Serengeti NP, 20% of cubs that emerge at >8 weeks survive to adulthood, but the short grass habitat here is likely particularly unfavourable for survival; mortality is lower everywhere else where data exist. Other post-emergence estimates include 38% (Phinda GR, South Africa), 43% (Nairobi NP, Kenya), 50% (Kruger NP) and 73% (S Kalahari). Adults occasionally killed by Lion, Leopard and Spotted Hyaena; males killed in territorial fights, and some adults killed in hunting accidents. Despite high genetic homogeneity, wild Cheetahs suffer little disease. LIFESPAN Maximum 13.5 (average 6.2) years for Serengeti females, 11 (average 5.3) for males; 21 in captivity.

Status and Threats

Reduced to approximately 7,100 adults and subadults in 33 known populations occupying 9% of the historic global range. More than 50% of Cheetahs occur in a single transboundary population across 6 southern African countries. Most of the range (77%) is non-protected where declines are poorly understood, but likely sufficient to justify upgrading the Cheetah’s status to Endangered. Widely persecuted by livestock farmers despite causing relatively minor damage, and profoundly affected by reduction of prey in pastoral areas. Human hunting of prey is critical in the Sahel, N Africa and Iran, where Cheetahs are naturally very rare. Limited hunting for skins occurs, e.g. Sahel and NE Africa, where there is also a significant trade in live cubs and adults, mainly to the Arabian Peninsula. High genetic homogeneity has had little impact on wild populations. CITES Appendix I, permitting trade of approximately 200 hunting trophies (Namibia and Zimbabwe) and live animals; Red List VU (global), CR (Asia), population trend Decreasing.

Plate 14

SNOW LEOPARD
Panthera uncia

Plate 15

OUNCE

HB 86–117cm, 104–125cm; T 78–105cm; SH to 60cm; W 21–53kg, 25–55kg

Dark cream to smoky grey with large, dark grey or black open blotches, and smaller solid markings on the head and legs. Muscular tail proportionally the longest of any felid, used for balance during hunts and wrapped around the body to insulate it against extreme cold. The species is unmistakable, but C Asian Leopards in long, pale winter coat are sometimes mistaken for it (chiefly in fur markets). Distribution and Habitat Found in 12 countries in C Asia throughout the world’s highest mountain ranges, including the Altai, Himalayas, Karakoram, Hindu Kush, Pamirs and Tien Shan. 2017 surveys in N Myanmar failed to confirm its presence. Uniquely adapted to high and steep rugged terrain, and copes well with deep snow. Also uses meadows, steppes, wide valleys and open montane desert, but mainly to move between rocky habitats. Occurs mostly at 3,000–5,500m except at northern range limits, where it is found at 900–2,400m, e.g. in Mongolia’s Gobi. Feeding Ecology Dependent on mountain ungulates, especially Asiatic (Siberian) Ibex, Blue Sheep and Argali, which are the main prey in much of its range. Other prey includes Markhor, Urial, Himalayan Tahr, musk deer, and occasionally gazelles, and juveniles of Eurasian Wild Boar, Wild Yak and Asiatic Wild Ass. Large kills are supplemented by small prey, especially during the summer, when herbivores disperse to higher elevations. Marmots are the most important of smaller prey; also opportunistically takes rabbits, hares, pikas, and game birds such as Tibetan Snowcock and partridges. Kills domestic animals, especially sheep and goats, but also cattle, yaks, camels, horses and dogs; depredation can reach significant levels locally and seasonally when wild prey is scarce. No recorded fatal attacks on humans. Stalks in typical felid fashion before rushing prey at close range; superbly agile over extraordinarily steep and rugged terrain. Large prey is usually killed by a suffocating throat bite. Eats carrion and has been observed displacing Dholes from carcasses (Hemis NP, India). Social and Spatial Behaviour Solitary. Maintains stable ranges that are regularly marked, but extent of territorial defence is unknown. Ranges are very large; published estimates from ground-based radio-telemetry are probably gross underestimates, e.g. a Mongolian female’s calculated range went from 58km2 to at least 1,590km2 (and possibly >4,500km2) when fitted with a satellite collar. Range size 87.2–193.2km2 (s) and 114.3–394.1km2 (s; Tost Mountains, Mongolia), assuming Snow Leopards closely follow the edges of mountain ranges; increases to 202.3–548.5km2 (s) and 264.9–1,283km2 (s) if associated steppe areas are included, but Snow Leopards mostly avoid steppe habitat. Species is extremely difficult to count, but density estimates from camera-trapping vary from 0.15/100km2 (Sarychat, Kyrgyzstan) to 4.5/100km2 in prey-rich habitat (Hemis NP, India). Reproduction and Demography Poorly known from the wild; likely to be a seasonal breeder given that it experiences extreme winters. Calling and scent-marking peaks January–March, which would coincide with spring births; captive births peak May. Gestation 90–105 days. Litter size averages 2–3 cubs, exceptionally to 5. Weaning at 2–3 months. Age at independence 18–24 months (2 individuals; Mongolia). MORTALITY Estimated at 17% (adults) to 23% (subadults; Tost Mountains, Mongolia). Causes are poorly known; winter starvation, especially of young animals, is likely to be an important factor. LIFESPAN 20 years in captivity. Status and Threats Somewhat insulated from human activities given that it lives in such remote, inhospitable areas, but it is naturally rare, and human populations and their livestock are increasing in its habitat. Widely killed for livestock depredation, often compounded by prey loss from human hunting and competition with livestock. Furs and especially body parts have commercial value, chiefly in China. CITES Appendix I; Red List VU, population trend Decreasing.

MAINLAND CLOUDED LEOPARD
Neofelis nebulosa
SUNDA CLOUDED LEOPARD
Neofelis diardi

Plate 15

HB 68.6–94cm, 81.3–108cm; T 60–92cm; W 10–11.5kg, 17.7–25kg

Classified as a single species until 2006, but now separated into 2 species: the insular Sunda Clouded Leopard; and the continental Mainland Clouded Leopard. Sunda Clouded Leopards are generally darker, with smaller cloud markings. Clouded leopards display an exceptionally large gape and elongated canine teeth, the reasons for which are unclear. Distribution and Habitat Mainland Clouded Leopard: continental Indochina. Sunda Clouded Leopard: Borneo and Sumatra. Both species are closely associated with dense evergreen tropical forest, but they also occur in secondary and logged forests, swamp forest, dry woodland and mangroves. Radio-collared individuals used grassland patches in open forest mosaics for hunting. Feeding Ecology Poorly known; confirmed prey includes primates from slow lorises to adult male Proboscis Monkey, hog deer, muntjacs, Bornean Bearded Pig, Malayan Pangolin, Asiatic Brush-tailed Porcupine, small rodents, civets and birds. Predation on orangutans by Sunda Clouded Leopard is unconfirmed. Unambiguous kills of deer and pigs revealed a deep killing bite to the nape, an unusual technique for large prey and perhaps related to the unique dentition. Recorded occasionally killing small livestock and poultry. Clouded leopards probably hunt mostly on the ground, but they are highly agile and arboreal; there are 4 published observations of clouded leopards attacking Proboscis Monkeys in trees. A female (Danum Valley, Borneo) cached an adult male Maroon Leaf Langur in a tree, where she fed on the carcass; it is unknown how frequently hauling occurs. Social and Spatial Behaviour Solitary. Only 8 animals have ever been radio-collared, in Borneo, Nepal and Thailand. Both species likely follow typical felid pattern of overlapping territories with exclusive core areas. Limited data suggest sexes have similar territory sizes: 34–40km2 (s) and 35.5–43.5km2 (s). Density estimates: 0.8−2.6/100km2 for Sunda Clouded Leopard (Sabah; Sumatra), and 0.6−3/100km2 (Htamanthi WS, Myanmar) to 5.1/100km2 (Dampa TR, India) for Mainland Clouded Leopard. Reproduction and Demography Probably aseasonal. Gestation 85–95 days (rarely to 109 days). Litters average 2–3 (range 1–5). Weaning begins at around 7–10 weeks. Sexual maturity at 20–30 months. MORTALITY Humans are responsible for most deaths in studied areas; otherwise mortality is unknown. LIFESPAN 17 years in captivity. Status and Threats Often assumed to be more resilient than Tigers and Leopards, but surveyed populations do not reach comparably high densities and both species are closely associated with forest habitat, which is under intense pressure for rubber and palm plantations or agriculture. Illegally hunted for skins, bones and meat. Both species: CITES Appendix I; Red List VU, population trend Decreasing.

Plate 15

TIGER
Panthera tigris

Plate 16

HB 146–177cm, 189–300cm; T 72–109cm; SH 80–110cm; W 75–177kg, 100–261kg

Background colour varies from pale yellow to rich red, with white or cream underparts. Generally darker and more richly striped in tropical S Asia, and paler and more lightly striped in temperate areas. White Tigers are not albino, and arise from a recessive mutation that produces blue eyes and chocolate-coloured stripes on a white background. There is only 1 record from the wild (Madhya Pradesh, India) since 1951, a male cub from which all captive white Tigers are descended (and, hence, are extremely inbred). An intermediate form, called ‘golden tabby’ or ‘strawberry’, is known only from captivity. True melanism is unknown, although pseudo-melanism occurs rarely, with extensive coalesced striping producing an almost entirely black appearance. Size varies very widely: Sumatran Tigers are the smallest, with males up to 140kg, while individuals from the Indian subcontinent and Russia are the largest. Largest wild male on record, from Nepal, weighed 261kg (up to 325kg recorded from captivity). Traditionally classified into 9 subspecies (three – Bali, Javan and Caspian – are now extinct) based mainly on morphology; incorporating comprehensive data, including genetic analyses, this was recently revised to 2 subspecies, the insular P. t. sondaica, now present only in Sumatra, and the continental P. t. tigris. Based on modest differences insufficient for subspecific divisions, continental Tigers are further grouped into a northern clade (Amur Tigers and the extinct Caspian Tiger) and a southern clade (all other mainland populations; Bengal Tiger).

Distribution and Habitat

Restricted to 7% of its historic range, from W India to the Himalayas, through Indochina to Malaysia, Sumatra, the Russian Far East and extreme NE China. Extinct in Bali (1940s), C Asia (1968), Java (1980s), SC China (by ~2000) and Cambodia/Vietnam/Laos (by ~2015). Occurs mainly in various temperate and tropical forests, forest–grassland mosaics and associated dense cover such as Terai grassland, thickets, scrub, marshes, mangroves and reed beds. Reaches the highest densities on the Indian subcontinent in dry and mesic forests, and the Terai. Typically avoids human-modified habitats such as agricultural land, palm plantations and monocultures. Occurs exceptionally up to 4,201m (Himalayas, Bhutan), but typically below 2,000m.

Feeding Ecology

Adults can kill almost anything they encounter, with the exception of adult rhinos and elephants, but the diet is dominated by various deer species and wild pigs. Typically focuses on ≤5 species of locally common prey such as Sambar, Red Deer, Chital, hog deer, Sika Deer, muntjacs and Wild Boar. Capable of killing adult Gaur and Water Buffalo weighing up to 1,000kg, but most kills of these species are subadults and juveniles. Smaller prey taken relatively often includes primates, porcupines, small carnivores and hares. Readily kills other predators, although they are not always eaten; Leopard, Dhole, Brown Bear and Asiatic Black Bear are recorded prey. Preys on livestock, mainly when untended in forest. Amur Tigers regularly kill domestic dogs, particularly when these are accompanying hunters in forest, and during severe winters that force the Tigers into villages. Tigers probably kill more people than any other large carnivore (mainly in India), in part because human populations in Asia are so dense and often utilise Tiger habitat intensively; true ‘man-eaters’ that focus on humans as prey are rare. Hunting is mainly nocturno-crepuscular. Scavenges and appropriates kills of other carnivores.

Social and Spatial Behaviour

Solitary and territorial. Adults establish exclusive territories where possible, but total exclusivity is rare. Territorial overlap is least in high-density populations with abundant prey and small ranges, e.g. Nepal, India. Territorial fights are rare, but sometimes fatal when they occur, more frequently in males than females. Recorded territory size varies from 10km2 (s; Chitwan NP, Nepal) to 2,058km2 (s; Russia). Range size for Terai and productive forest in Nepal or India 10–51km2 (s) and 24–243km2 (s), compared with 181–761km2 (s) and 434–2,058km2 (s) in Russia. Females mostly settle near their natal range, while males disperse more widely. Dispersal distances at Chitwan NP average 9.7km for females (maximum 33km) and 33km for males (maximum 65km), compared with 14km for females (maximum 72km) and 103km for males (maximum 195km) in Russia. Densities of populations, even in high-quality habitat, are often depressed due to human hunting of prey (even if Tigers are not hunted). Density estimates vary from 0.2–2.6/100km2 in lowland tropical forest where poaching is prevalent (Laos; Malaysia; Myanmar; Sumatra) and 0.5–1.4/100km2 (temperate forest, Russia), to 11.5–19/100km2 in well-protected forest and Terai (India).

Reproduction and Demography

Aseasonal over most of the range; more than 50% of Amur Tiger cubs are born late summer, August–October, and winter births are rare. Oestrus 2–5 days; gestation 95–107 days, averaging 103–105. Litter size 2–5, averaging 2–3. Weaning at around 3–5 months. Inter-litter interval 20–24 months, averaging 21 months. Cubs independent at 17–24 months. Females often inherit part of their mother’s range, while males disperse more widely. Sexual maturity at 2.5–3 years (both sexes): earliest breeding 3 years for females (average 3.4 years in Chitwan NP) and 3.4 years for males (average 4.8 years, Chitwan NP). Reproduction by females is possible until at least age 15.5. MORTALITY 34% (Chitwan NP) to 41–47% (Russia) of cubs die in their first year, most related to anthropogenic causes and infanticide. Estimates of adult mortality include 23% (both sexes combined; Nagarahole NP, India) and 19% (s) to 37% (s; Russia). Humans are the main cause of death for most populations, and adults also die in territorial fights. Accidents (e.g. a male fell through a frozen river in Russia) and disease occur, but are uncommon. LIFESPAN 16 years for females and 12 for males in the wild; 26 in captivity.

Status and Threats

The most endangered large cat, having suffered a calamitous decline in the twentieth century that continues today, with approximately 3,900 individuals in 9 range countries. An estimated 70% of the world’s Tigers (and most breeding females) now live in around 0.5% of their historic range. Combined with loss of habitat to forestry, commercial palm plantations and agriculture, Tigers are particularly threatened by intense illegal hunting to supply the traditional Asian medicinal trade. This is compounded by widespread hunting of their prey to feed a massive demand for bushmeat, especially in Southeast Asia. Given strong protection, Tiger populations recover rapidly, e.g. densities increased from 0.8/100km2 to 1.4/100km2 between 2013 and 2016 in Parsa NP, Nepal, but unfortunately there are few areas where sufficient protection is taking place. CITES Appendix I; Red List EN (globally), CR (China, Russia, Sumatra), population trend Decreasing.

Plate 16

LION
Panthera leo

Plate 17

HB 158–192cm, 172–250cm; T 60–100cm; SH 100–128cm; W 110–168kg, 150–272kg

Typically tawny or sandy with pale underparts, but varying from ash grey or cream to ginger and (rarely) dark brown. Backs of the ears and tail tip are a contrasting black or dark brown. White Lions from Kruger NP region, South Africa, are leucistic (not albino), with pigmented eyes, nose and pads; they can be born to normally coloured parents. Mane colour ranges from platinum to black, and the length is highly variable. ‘Maneless’ males occur most often where it is extremely hot, e.g. Tsavo NP, Kenya, and in most Sahelian populations. Cubs are born with dark brown rosettes that fade with age, retained as vestigial spotting in some adults. Mane growth begins at 6–8 months. Recent molecular studies show a clear separation into a northern subspecies, P. l. leo (W and C Africa, and India), and southern subspecies, P. l. melanochaita (E and southern Africa). Northern Lions are 10–20% smaller than E and southern African individuals, and males typically have reduced manes; Indian Lions often have a distinctive belly fold (occasionally present in African Lions).

Distribution and Habitat

Patchy distribution south of the Sahara, chiefly in and around protected areas, with the largest populations in E and southern Africa. Outside Africa, there is only 1 population, in Gujarat, India. Naturally absent only from true desert and equatorial rainforest. Optimum habitat is mesic open woodland and grassland savannah. Inhabits dry deciduous teak forest in India. Recorded to 4,200–4,300m (Bale Mountains, Ethiopia; Mt Kilimanjaro, Tanzania).

Feeding Ecology

Opportunistic, killing virtually everything it encounters, but prefers large herbivores weighing 60–550kg; cannot persist without large prey. In any given population, the diet is dominated by 3–5 ungulates such as wildebeest, zebra, buffalo, giraffe, Gemsbok, Impala, Nyala, Kob, Thomson’s Gazelle, Chital, Sambar and Warthog. Smallest preferred prey species documented is Springbok (27–48kg), in Etosha NP, Namibia. Smaller prey, especially Warthog, often dominates during the lean dry season where large prey is migratory, e.g. Chobe NP, Botswana and Serengeti NP, Tanzania. Only healthy mature elephants are invulnerable to Lion predation; large prides kill young or unwell adults, as well as adult rhinos and Hippopotamuses. Kills untended livestock and occasionally preys on humans; isolated pockets of persistent man-eating still occur, e.g. SE Tanzania and N Mozambique. Hunting is mainly nocturno-crepuscular. Although females make most kills, males are capable hunters that increase the hunting success of very large prey, e.g. buffalo and giraffe. Success estimates include 15% (Etosha NP), 23% (Serengeti NP) and 38.5% (Kalahari). Readily scavenges, comprising 5.5% of intake in Etosha NP to almost 40% in Serengeti NP. Frequently appropriates kills from other carnivores.

Social and Spatial Behaviour

The only communally living cat. Prides comprise 1–20 (usually 3–6) related lionesses, their offspring and 1–9 (usually 2–4) immigrant males unrelated to breeding females. Pride size is correlated with prey biomass, smaller in arid areas and exceptionally reaching 45–50 under ideal conditions. Female membership of the pride is mostly stable, but the entire pride is together rarely: small subgroups come and go within pride range in a continuous ‘fission–fusion’ pattern. Females defend their range against other prides and strange males. Coalition males are usually related to each other, but smaller coalitions (2–3) often include unrelated members. Females generally stay with the pride for life, but occasionally disperse following a male takeover or to avoid mating with male relatives. Young males are evicted or leave at 25–48 months, entering a nomadic stage lasting 2–3 years before attempting to acquire their own pride. Coalition members are highly cooperative and remain together for life, defending their territory and females from male intruders. Following a takeover, new males usually kill or evict all unrelated cubs younger than 12–18 months to hasten the lionesses’ return to oestrus. Coalition tenure is generally 2–4 years. Serengeti pride territories average 65km2 (woodland) to 184km2 (grassland), reaching a maximum of 500km2. Arid areas produce much larger ranges: to 2,800km2 (Kalahari), 1,055–1,745km2 (Kaudom GR, Namibia) and 2,721–6,542km2 (Kunene, NW Namibia). Two Kunene male coalitions (possibly nomads) had ranges of 13,365–17,221km2. Density estimates: 0.05–0.62/100km2 (Kunene), 1.5–2/100km2 (Kalahari), 6–12/100km2 (Kruger NP), 12–14/100km2 (Gir PA, India), to 38/100km2 (Lake Manyara NP, Tanzania).

Reproduction and Demography

Aseasonal, although births often peak with seasonal birth events of ungulates. Oestrus averages 4–5 days; gestation 98–115 days (mean 110). Litter size typically 2–4, up to 7. Lionesses often give birth synchronously and communally care for cubs; females suckle all cubs, but carry only their own. Weaning begins at around 6–8 weeks, but suckling may continue to 8 months. Cubs can hunt independently at around 18 months, but rarely disperse before 2 years. Inter-litter interval 20–24 months. Lionesses can conceive at 30–36 months, but typically first give birth at around 42–48 months and cease reproducing after age 15. Males are sexually mature at 26–28 months, but rarely breed before 4 years. MORTALITY 16% (Kruger NP, with abundant resident prey) to 63% (Serengeti NP, with mainly migratory prey) of cubs die in their first year, mostly from infanticide, predation and starvation; mortality in the second year typically drops significantly, from 10% (Kruger NP) to 20% (Serengeti NP). Apart from human-caused mortality, adult Lions die mostly in fights with other Lions (especially males), from injuries while hunting large prey and from starvation when old or debilitated. Disease is uncommon, but episodes are occasionally severe; more than 1,000 Lions (40% of the population) died during the 1993–94 canine distemper outbreak, Serengeti NP. LIFESPAN 19 years for females, 16 (but rarely over 12) for males in the wild; 30 in captivity.

Status and Threats

Conservation-dependent, inhabiting <16% of historic African range, with 1 isolated population of 400 in India. Globally, the total population declined by 43% in 1993–2014; numbers in 4 southern African countries (Botswana, Namibia, South Africa and Zimbabwe) and India collectively increased by 12%, while populations in the rest of the range declined by 60%. Excluding southern Africa/India, the Lion’s status is Endangered. Eradication by humans, coupled with combined loss of habitat and prey from agriculture and livestock herding, has driven declines. Intense persecution by herders (mainly outside protected areas) and poaching for bushmeat (mostly inside protected areas) are the primary ongoing threats. Highly vulnerable to poisoned baits, and to snares set for bushmeat; a majority of protected populations are significantly depleted due to poaching, mainly of prey but also ancillary poaching of Lions. Legal sport hunting takes place in 10 African countries. Lions are very sensitive to overharvest, and hunting triggers population declines under excessive quotas or where they are already suffering anthropogenic mortality, such as from snares. Excluding canned hunts of captive-bred lions in South Africa, approximately 220–240 wild Lions (mainly males) are hunted annually. CITES Appendix I – Asia, Appendix II – Africa; Red List VU (global), EN (India), CR (W Africa), population trend Decreasing.

Plate 17

LEOPARD
Panthera pardus

Plate 18

PANTHER

HB 95–123cm, 91–191cm; T 51–101cm; SH 55–82cm; W 17–42kg, 20–90kg

Background colour varies from pale cream, through various shades of orange, to dark rufous-brown with white underparts, covered with rosettes, each a cluster of small black spots around a normally unspotted centre that is darker than the body colour. Forest Leopards tend to be dark, while those in arid areas are pale. Melanistic individuals (‘black panthers’) occur mainly in humid lowland and upland forests; they are most common in tropical Southeast Asia, e.g. outnumbering spotted Leopards in Malaysia, and are rarely recorded in Africa. Size varies widely, correlated with changes in climate and prey availability. The smallest Leopards, from arid mountainous areas in the Middle East, are about half the weight of African woodland-savannah individuals. Leopards from an isolated population in coastal Cape Mountains, South Africa, are also small, averaging 21kg (s) to 31kg (s). The largest Leopards are recorded from E and southern African woodland and N Iran.

Distribution and Habitat

Widely distributed in southern, E and C Africa, significantly reduced in W Africa and most of Asia, and relict in N Africa, the Middle East and Russia. Very wide habitat tolerance, ranging from Russian boreal forests with winter lows of –30°C, to desert with summer highs of 70°C. Reaches the highest densities in mesic woodland, grassland savannah and forest, and fairly common in mountains, scrub and semi-desert. Absent from open interiors of true desert, but occupies watercourses and rocky massifs in very arid areas. Tolerates human-modified landscapes provided cover and prey are available, e.g. coffee plantations and sugar-cane fields. Recorded exceptionally to 5,638m (Mt Kilimanjaro, Tanzania).

Feeding Ecology

Extremely catholic, with prey ranging from arthropods to adult male Elands (maximum weight 900kg), but prefers medium-sized ungulates weighing 15–80kg. Typical prey includes Steenbok, duikers, Impala, gazelles, Nyala, muntjacs, Chital, Siberian Roe Deer, Bushpig and Warthog, and young individuals of larger animals such as Gaur, wildebeest, oryx, hartebeest, Greater Kudu, Sambar and Wild Boar. Additionally, primates, hares, rodents, small carnivores and large birds are often important. Preys on livestock, occasionally entering corrals and settlements, and readily kills domestic dogs. Sometimes preys on humans. A consummate stalk-ambush hunter, approaching prey to as close as 4–5m before a final explosive rush. Hunting is mainly nocturno-crepuscular, and most daylight hunts are unsuccessful. Hunting success estimates include 15.6% (Kalahari), 20.1% (Phinda GR, South Africa) and 38.1% (NE Namibia). Kalahari Leopards average 111 (s) to 243 (s) kills annually, with females making more smaller kills. Leopards hoist carcasses weighing up to 91kg into trees to avoid kleptoparasitism, primarily from Spotted Hyaenas, and occasionally also cache in caves, burrows and kopjes. They typically pluck fur before feeding, usually starting at the underbelly or hind legs. They readily scavenge.

Social and Spatial Behaviour

Solitary and territorial. Adults defend a core area against same-sex conspecifics, but tolerate considerable overlap at the edges, with mutual avoidance and alternating use of shared areas. Territorial fights are uncommon, but may result in fatalities in both females and males. Males associate with familiar females and cubs for as long as 24 hours, but never form permanent family groups. Recorded territory size 5.6km2 (s; Tsavo NP, Kenya) to 2,750.1km2 (s; Kalahari). Mean range size for mesic woodland, savannah and rainforest across the species’ distribution averages 9–27km2 (s) and 52–136km2 (s). Ranges are much larger in arid habitats, averaging 188.4km2 (s) and 451.2km2 (s) in N Namibia, and 488.7km2 (s) and 2,321.5km2 (s) in Kalahari. A collared male in arid rocky habitat, C Iran, used 626km2 in 10 months. Density estimates: 0.5/100km2 (Etosha NP, Namibia), 1–1.4/100km2 (Primorsky Krai, Russia), 1.3/100km2 (Kalahari), 4.6–12/100km2 (Gabon rainforest) and 11.1/100km2 (Phinda and Mkhuze GRs, South Africa), to 12/100km2 (Sabi Sands GR, South Africa). In African woodland savannah, average density under protection is almost 5 times as high as outside protected areas.

Reproduction and Demography

Poorly known in its northern range (China, N Korea, Russia), where extreme winters might give rise to seasonality; otherwise aseasonal. Oestrus 7–14 days; gestation 90–106 days. Litter size normally 1–4, rarely to 6 (captivity). Weaning begins at around 8–10 weeks and suckling typically ceases before 4 months. Inter-litter interval averages 16–25 months. Cubs independent at 12–18 months; earliest age at which cubs survive independently is 7 months. Female dispersers often inherit part of their mother’s range, while males disperse more widely. Both sexes are sexually mature at 24–28 months; females first give birth at 30–36 months, and can reproduce to 16 years (19 in captivity). Males first breed at around 42–48 months. MORTALITY 50% (Kruger NP) to 62% (Sabi Sands GR) of cubs die in their first year; where well-studied (savannah Africa), most deaths are by male Leopards, followed by Lions, less so Spotted Hyaenas. Estimates of adult mortality include 18.5% (Kruger NP) to 25.2% (Phinda GR, South Africa). Aside from deaths attributable to humans, adults are killed primarily in territorial fights and by other predators, chiefly Lions and occasionally Tigers, Spotted Hyaenas, African Wild Dog and Dhole packs, baboon troops (rarely) and large crocodiles. Hunting accidents and deaths from disease are uncommon. LIFESPAN 19 years for females, 14 for males in the wild; 23 in captivity.

Status and Threats

Surprisingly tolerant of human activity and persists where other large carnivores cannot. Even so, extirpated from approximately 75% of global historic range (48–67% of African range and 83–87% of Eurasian range). Arabian (P. p. nimr), Amur (P. p. orientalis), Indochinese (P. p. delacouri) and North Chinese (P. p. japonensis) subspecies each inhabit <5% of historic range and should all be regarded as Critically Endangered. Loss of habitat and prey, closely followed by intense persecution in livestock areas, are the chief threats. Heavily hunted in S Asia for skins and parts supplying the Chinese medicinal trade, and killed for skins, canines and claws in W and C Africa. Bushmeat hunting, especially in tropical forest, competes directly for principal prey species and may drive extinctions even in intact forest. Legal sport hunting takes place in 9 African countries; Namibia (78–123 killed annually, 2012–15), Tanzania (163–205 killed) and Zimbabwe (136–163 killed) account for the highest offtake. Excessive quotas contribute to population declines, leading some countries to ban (Botswana) or suspend (South Africa, Zambia) hunting. CITES Appendix I; Red List VU (global), EN (Sri Lanka, C Asia), CR (Java, Middle East, Russia), population trend Decreasing.

Plate 18

JAGUAR
Panthera onca

Plate 19

HB 116–219cm, 110.5–270cm; T 44–80cm; SH 68–75cm; W 36–100kg, 36–158kg

The world’s third largest cat. Background colour varies from pale yellow through ginger to rufous-golden brown, with white or cream underparts. Large block-like markings or rosettes usually enclose smaller black spots (usually lacking in the similar Leopard). Melanism occurs, with the same pattern of markings apparent in oblique light; black individuals are most common in humid lowland rainforest. Size varies very widely; the smallest Jaguars occur in Central America and are about half the size of the largest individuals from wet woodland savannah habitats of Brazil and Venezuela.

Distribution and Habitat

N Mexico to N Argentina. Widely distributed in much of N and C South America, more fragmented in Mexico, Central America, E Brazil, Argentina and S Bolivia. Resident breeding populations no longer occur in the USA, but individuals intermittently appear in Arizona and New Mexico from N Mexico. Broad habitat tolerance; while capable of occupying dry open savannah or desert habitats, the species is more commonly associated with the dense cover of tropical and subtropical lowland forest, typically below 2,000m (exceptionally to 2,700m in the Andes, 3,000m in Mexico). Strongly associated with water, and thrives in well-watered habitats, including flooded savannah (Brazilian Pantanal and Colombian/Venezuelan Llanos), swamps, riverine thicket and scrub, and mangroves. Excellent swimmer, capable of crossing large rivers >2km wide.

Feeding Ecology

Diverse diet, with at least 85 recorded prey species. Like all large cats, focuses on common large-bodied prey, but the natural absence of herds of large deer and wild cattle in South America results in the hunting of smaller animals more often than is the case for other big cats. Capybara and Collared and White-lipped Peccaries are frequently the most important prey species where they occur. Reptiles form a larger part of the diet than in any other large cat, especially caimans, as well as iguanas, freshwater turtles and tortoises, nesting marine turtles and large boas such as anacondas. Can subsist on abundant small prey such as armadillos, pacas, brocket deer and/or coatis. Other relatively common prey includes White-tailed Deer, agoutis, marsupials and sloths. Occasionally kills very large prey, including tapirs and Marsh Deer; Jaguars in the Brazilian Pantanal are recorded killing Amazon River Dolphins as they fish in shallow water. Readily kills domestic livestock; in ranching-dominated habitats such as the Pantanal and Llanos, introduced cattle form the major prey species. Less often, takes domestic pigs and dogs from villages. Almost never hunts humans; most recorded attacks result from extreme provocation, for example during Jaguar hunts, and verified unprovoked attacks are extremely rare. Jaguars have proportionally the strongest bite of all large cats, and kill either by biting with massive force at the back of the skull or with a typically feline suffocating throat bite. Hunting is mainly nocturno-crepuscular and terrestrial, although Jaguars readily hunt prey, e.g. Capybara and caimans, in water. Often scavenges, including from cattle carcasses; the deaths of these cattle are often erroneously blamed on the cat.

Social and Spatial Behaviour

Solitary and territorial, but exclusive range use appears limited to small core areas; overlap between adults in some populations is extensive, possibly due to marked seasonal changes in the distribution of water and hence prey. Pantanal females establish largely exclusive ranges during the wet season, but overlap considerably in the dry season, while males overlap extensively in both. Similarly, Cockscomb Basin WS (Belize) males overlap extensively. Adults engage in typically territorial behaviours such as roaring, scrapes and urine-marking, perhaps serving to foster avoidance rather than demarcate exclusivity. Aggressive interaction between adults appears to be rare, although there are records of fatal fights. Range size estimates include 28–40km2 (s) in Belize; 38km2 (average, s) to 63km2 (average, s) in S Pantanal, Brazil; 47–83km2 (s) to 93–108km2 (s) in Venezuela, and 31–98km2 (s) to 73–268km2 (s) in N Pantanal, Brazil. A male in arid lowland desert and pine–oak woodland in Arizona used at least 1,359km2 in 2004–07. Home range sizes in inundated habitat, e.g. Pantanal, often contract in the wet season, when flooded areas limit space available to prey. Density estimates: 1.1/100km2 (Iguaçu NP, Brazil), 2.5/100km2 (Atlantic forest, Brazil), 3.5/100km2 (Corcovado NP, Costa Rica) and 8.8/100km2 (Cockscomb Basin WS), to 6–11/100km2 (Pantanal).

Reproduction and Demography

Aseasonal. Oestrus 6–17 days; gestation 91–111 days (average around 101–105 days). Litter size 1–4, averaging 2 (captivity). Weaning begins at around 10 weeks, and suckling typically ceases by 4–5 months. Cubs independent at 16–24 months. Dispersal poorly known, but appears to be typically feline, in which females settle close to their natal range while males disperse more widely. Both sexes are sexually mature at 24–30 months; females first give birth at 3–3.5 years and can reproduce to 15 years. It is not known when wild males first breed. MORTALITY Rates are poorly known. Adult Jaguars have no predators and are killed principally by humans and rarely by other Jaguars. Humans remove very significant numbers from some populations, e.g. an estimated 230 Jaguars killed in 1989–2014 by people living adjacent to Cerro Hoya and Darién NPs, Panama; and 539 Jaguars killed in 2001–15 in rural localities across Venezuela. Predators of cubs are poorly known; infanticide by male Jaguars is recorded, and there is a record of an adult female killing an unrelated cub. LIFESPAN Poorly known from the wild, but unlikely to exceed 15–16 years; 22 in captivity.

Status and Threats

Extirpated from an estimated 49% of its historic range and extinct in El Salvador, Uruguay and the USA. Despite this, much of its remaining range is potentially still continuous, in part because the massive forested basins of South America have remained mostly inaccessible until recently. Habitat conversion for forestry, livestock and agriculture is the main threat, combined with intense persecution from ranchers and pastoralists in livestock areas, despite the fact that many cattle losses blamed on Jaguar predation occur from other factors. Widespread and intensive hunting of prey is likely to impact Jaguar populations in many areas, and emerging evidence indicates that the species is hunted to supply the Chinese medicinal trade, although impacts of both are poorly quantified. Sport hunting is illegal, but poorly enforced, in all range countries. CITES Appendix I; Red List NT, population trend Decreasing.

Plate 19