Cheetah

Acinonyx jubatus (Schreber, 1775)

IUCN RED LIST (2008):

Vulnerable (global)

Critically Endangered (Asia)

Critically Endangered (heckisubspecies)

Head-body length ♀ 105−140cm, ♀ 108−152cm

Tail 60−89cm

Shoulder height ♂ 67−89cm

Weight ♀ 21−51kg, ♂ 29−64kg

Taxonomy and phylogeny

The Cheetah is the sole species in the genus Acinonyx. With its closest living relatives, the Puma and Jaguarundi, it is part of the Puma lineage, which diverged around 6.7 million years ago. The oldest Cheetah fossils are 3–3.5 million years old from southern and East Africa.

Four African and one Asian subspecies are currently described. The Asiatic cheetah A. j. venaticus is the most distinct subspecies, with genetic differences indicating isolation from African populations for an estimated 32,000–67,000 years. Cheetahs in north-eastern Africa (traditionally A. j. soemmeringii, the Central African Cheetah, Somalia to eastern Niger) are genetically distinct from other African populations, though they are likely contiguous with the West African cheetah (traditionally A. j. hecki, western Niger to Senegal).

Further molecular research may combine them as one subspecies. Similarly, African Cheetahs north of the Sahara were formerly considered A. j. venaticus but a western Egypt sample shows they are not closely related to Iranian cheetahs, and are likely to cluster with soemmeringii/hecki. Cheetahs from southern Africa (A. j. jubatus) and East Africa (A. j. raineyii) are closely related, with relatively minor genetic differentiation, though they are presently considered two subspecies.

Description

The Cheetah is the only felid adapted for prolonged high-speed pursuits of prey, with an unmistakable, greyhound-like build unlike any other cat. It has a tall, slim frame with long legs, a narrow, deep chest, narrow waist and long, tubular tail. The head is small and rounded, with a short muzzle and small ears. Cheetah claws are dog-like and they lack the fleshy, protective claw-sheaths present in other cats, but, contrary to popular belief, the claws are partially protractile. Claws appear in the tracks except for the sharp, strongly curved dew claw that is used for prey capture. Southern African cheetahs tend to be largest, closely followed by those from East Africa. The smallest Cheetahs (based on very few measured specimens) are from the Sahara, north-eastern Africa and Iran.

The Cheetah is typically light tawny to yellow-blonde fading to creamy-white underparts. About 2,000 solid black round or oval spots cover the body, interspersed sporadically with small, often indistinct black dabs. The pattern these markings make is unique to each individual. The Cheetah’s lachrymal or ‘tear’ streaks are not found in any other felid and their function, if any, is unclear – they may help to reduce glare for diurnal hunting and enhance defensive or aggressive facial expressions. Saharan Cheetahs tend to have distinctive narrow, dog-like faces and short, pale fur ranging from yellow-beige with chocolate-brown spots to near-white with faint cinnamon or ochre markings. The so-called King Cheetah was considered a separate subspecies A. j. rex when first described in 1927 but is merely a recessive colour variant which may be born to normally spotted parents. In the wild, the king variant appears regularly only in northern South Africa (including Kruger National Park), southern Zimbabwe and south-eastern Botswana. A single king skin was confiscated from a poacher in Burkina Faso, possibly a trade skin. Melanism is extremely rare, with only two confirmed specimens, from Zimbabwe and Zambia. There are no confirmed records of true albino Cheetahs, but a yellow-cream-coloured Cheetah with dark freckling was photographed in Kenya’s Athi River region in 2010.

Uniquely among cats, Cheetah cubs are born with a fluffy, smoky-grey mantle that runs from the crown to the base of the tail. The mantle dwindles by four to six months to become a short mane on the shoulders that is inconspicuous in most adults except when raised in fear or aggression. The mantle is more prominent in Asiatic Cheetah adults, especially during winter when Iranian individuals develop long, silky coats. The function of the mantle is unclear. A popular theory suggests it mimics the irascible Honey Badger to deter predators. Although the similarity is striking, its value is dubious given the very high rates of predation on cubs in some populations. More likely, the mantle probably assists with thermoregulation and perhaps in camouflaging young, helpless cubs in long-grass dens.

Similar species The Cheetah is similar in overall size and colouration to the Leopard, but confusion is unlikely, except perhaps in fleeting sightings – the Cheetah’s simple spots and tear streaks are distinctive. Servals are often identified by local people as Cheetahs, though they are much smaller with a characteristic short tail.

The Cheetah is less sexually dimorphic than most large cat species. Nonetheless, adult males are significantly more heavily built than females, especially in the head, neck and forequarters.

One of the last remaining Asiatic Cheetahs, photographed by camera-trap in Naybandan Wildlife Refuge in Iran. This adult male shows the very short coat typical of Asiatic Cheetahs during the extremely hot Iranian summer.

Once considered a separate species, the King Cheetah is simply the equivalent of a tabby-coloured Cheetah. A mutation in the same gene that produces tabby markings in domestic cats causes the spots to coalesce into blotches and stripes.

Distribution and habitat

The Cheetah is relatively widely distributed in southern and East Africa, rare to very rare in West and Central Africa and extinct in North Africa except southern Algeria and perhaps western Egypt. It is extinct in Asia except for approximately 50 animals in central Iran. Occasional reports since the 1970s from Afghanistan, Pakistan and Turkmenistan lack unequivocal evidence. A skin found in Mazar-e-Sharif in 2006 reportedly came from Samangan Province, northern-central Afghanistan – it is more likely, however, to have originated in Iran.

The Cheetah favours savanna woodland, grasslands and scrublands, reaching its highest densities in mesic mosaics of open woodland and grassland. It is more sparsely distributed in dense, humid woodlands such as Miombo, and does not occur in thick forest and rainforest. They are well adapted to arid savannas such as in the southern Kalahari, and they inhabit semi-desert and desert in Iran, the Namib and the Sahara where they are associated with watercourses and mountain ranges; they are transient in the driest areas of true desert. Cheetahs are recorded exceptionally to 3,500m (Mount Kenya, Kenya) but suitable habitat for Cheetahs typically dwindles at around 1,500m (Ethiopia) to 2,000m (Algeria). Iranian Cheetahs live above the winter snowline in desert massifs, the only place where the species routinely experiences winter snow.

Provided they survive, these young brothers will remain together as a male coalition once they become independent. The strong bonds they form as cubs will endure for life, with almost constant contact, cooperative behaviour and affectionate displays.

Feeding ecology

The Cheetah is famously the fastest terrestrial animal on Earth, best suited for hunting small to medium-sized antelopes weighing 20−60kg; gazelles or gazelle-analogues are preferred throughout the range. Typical primary prey species include Thomson’s Gazelle and Grant’s Gazelle (East Africa), Dama Gazelle and Dorcas Gazelle (Sahara), Springbok (arid southern savannas, e.g. Etosha National Park and the Kalahari), and Impala (southern and East African woodlands, e.g. Kruger National Park, Okavango, Serengeti woodlands). Smaller antelopes such as Steenbok and Common Duiker are important prey, for example in the southern Kalahari. Cheetahs are capable of taking large ungulates, which may constitute preferred prey in areas where they are the most abundant option, e.g. Nyala (adults 55−127kg) in mesic woodland, Phinda Game Reserve, South Africa, or where smaller prey is unavailable, e.g. Urial sheep (adults 36−66kg) and Wild Goat (adults 25−90kg) in Iran following the widespread extirpation of gazelles. Generally, juveniles are preferred prey over adults and females are preferred over males, especially when prey is large or dangerous, such as Warthogs. Male gazelles are often seasonally preferred where they have a defined breeding season as males compete with other males making them less vigilant and more vulnerable to Cheetah predation, e.g. Springboks in the Kalahari. Cheetahs, typically male coalitions, readily kill juveniles of large species including Blue Wildebeest, Burchell’s Zebra, Gemsbok and, rarely, African Buffalo and Giraffe. Male coalitions are also capable of taking large, dangerous prey that individual male and female Cheetahs usually avoid, for example adult Hartebeest, oryx and wildebeest.

Large rodents, such as Spring Hares, and large lagomorphs (Cape Hares, Scrub Hare) are important prey in some situations, for example for single Kalahari Cheetahs, for recently independent young adults in many populations, and in impoverished habitats, e.g. in Iran. Cheetahs sometimes kill ostriches, bustards and guineafowl, and rarely hedgehogs (Sahara), porcupines and small carnivores including mongooses and foxes. Cannibalism has been recorded by males following fatal, territorial clashes but is otherwise unknown. Cheetahs in the Kalahari reputedly eat tsamma melons for the water content but a comprehensive six-year observational study did not record this, and it is unlikely tsammas contribute much if anything to water intake. Cheetahs kill small, untended livestock, such as goats, sheep, calves and juvenile camels, but they are easily deterred by people or dogs. There is no record of a wild Cheetah killing a human.

In the majority of their range, Cheetahs are most active during the day, maximising visibility and reducing overlap with peak activity periods of Lions and Spotted Hyaenas. Most hunts occur within a few hours of sunrise and sunset, but often throughout the day during cooler periods and if females have cubs. Nocturnal activity is more common than widely believed. About 25 per cent of all activity by Okavango cheetahs occurs during the night, mostly around full moons; nocturnal hunting occurs but it is unclear if 25 per cent of activity translates to 25 per cent of hunting effort. Saharan Cheetahs are thought to hunt frequently at night, probably taking advantage of cooler temperatures and very open habitat. Cheetahs usually initiate a hunt by stalking in fairly typical felid fashion, semi-crouched and freezing when prey looks up, to within 50–75m of the target. They may, however, start trotting openly towards very vulnerable or oblivious prey, such as neonate antelopes, from 500 to 600m away. Sprints last up to 600m but typically less than 300m; chases average 173m for Cheetahs hunting in open savanna woodlands (Okavango, Botswana). A trained cheetah running behind a vehicle was timed at 105kmph, the fastest recorded speed – the top speed recorded for wild Botswanan Cheetahs is 93kmph. The Cheetah is probably capable of exceeding 105kmph, but adults hunting mainly Impalas in Botswanan savanna woodlands use moderate speeds, with a top speed averaging 54kmph. Cheetahs hunting gazelles in very open habitat probably average faster hunting speeds. The Cheetah can accelerate and decelerate faster than any terrestrial mammal. Cheetahs decelerate very rapidly in the final stage of the hunt which enhances manoeuvrability for making the kill, e.g. from 58kmph to 14kmph in three strides (Okavango, Botswana). Prey is tripped or bowled over with the front paw, or pulled off balance by hooking with the dew claw and yanking backwards while decelerating. Most prey is killed by suffocation, taking 2–10 minutes, but hares and other small prey are killed rapidly by a bite to the skull or nape.

Around 25–40 per cent of hunts are successful, with higher success rates for hunts of small and vulnerable prey: 87−93 per cent for hares, 86−100 per cent for juvenile gazelles (Serengeti National Park, Tanzania). Cheetahs rarely defend their kills, losing up to 13 per cent (Serengeti National Park), chiefly to Spotted Hyaenas and Lions, less often to Leopards, Wild Dogs, Grey Wolves (Iran), Brown Hyaenas and Striped Hyaenas. Groups of Black-backed Jackals and large flocks of vultures are recorded usurping Cheetahs from their kills but they probably leave fearing that larger carnivores will be attracted by the commotion. Cheetahs very rarely scavenge, presumably because of the danger of encountering larger carnivores, but male Cheetahs readily take kills from female Cheetahs. Cheetahs also seldom revisit previously fed-upon carcasses for the same reason, but mothers with cubs sometimes return to a kill after abandoning it overnight (Phinda).

Killing by suffocation is a safe technique for subduing potentially dangerous prey. This Cheetah killing a young Blesbok shows how cats further reduce the chance of injury from struggling prey by immobilising the horned head and positioning themselves away from thrashing hooves.

Social and spatial behaviour

The Cheetah has a fluid and complex socio-spatial system unique among felids, with greater sociality than all other cats except the Lion (and some feral domestic cat colonies). Female Cheetahs are asocial and non-territorial with large home ranges that are not defended. Encounters between females typically result in both actively avoiding contact or, rarely, resting together tolerantly for less than a day (often because they are related; see below). In contrast, males are social and often form lifelong coalitions of two to four individuals that compete aggressively with other males for access to females. Coalitions usually comprise brothers who remain together after dispersal. Pairs or singletons, however, often recruit an unrelated member; for example, 30 per cent of Serengeti coalitions contain a non-relative. Some males, particularly those without male siblings, may never belong to a coalition – around 40 per cent of Kalahari males are solitary. Males may or may not be territorial; because females are semi-nomadic, males typically establish territories only where it increases access to females, for example when female ranges are small or their movements are predictable. Serengeti males establish small territories often centred on areas of cover and where Thomson’s Gazelles may aggregate seasonally. Female Cheetahs follow gazelle migrations and are attracted to the aggregations. In areas with non-migratory prey (e.g. southern Kruger National Park, South Africa), female ranges tend to be smaller with less nomadic movements, making male territoriality profitable. Male territories in this scenario are a similar size to female ranges. Both coalitions and single males may be territorial, but coalitions are more successful at defending territories and therefore more likely to gain access to females. Serengeti coalitions are also healthier than singletons. Territorial males repel rival males in fights, which are sometimes fatal.

As an alternative to territorial residency, males may be non-territorial, semi-nomadic ‘floaters’, either because they are unable to defend a territory or possibly because it is an equally successful strategy for finding females. All male Cheetahs start out as floaters and some never become resident; those that do may switch between territoriality and floating, depending on the availability of prey (and therefore of females) and whether they belong to a coalition. Floaters make up 40 per cent of the Serengeti population. All central Namibian males appear to be floaters with very large ranges, perhaps because of low prey densities and high human mortality creating constant disturbance in the population.

Range size is among the largest recorded for felids. Where prey is migratory or scarce, female range size averages 833km2 (range 395−1,270km2; Serengeti National Park) to 2,160km2 (range 554−7,063km2; central Namibia). Female ranges are smaller in habitat with resident or abundant prey, e.g. 34−157km2 (woodland, Phinda Game Reserve) and 185−246km2 (woodland, Kruger National Park). Male territory size averages 33−42km2 (single males and coalitions, Serengeti) and 93km2 (range 57−161km2; coalitions, Phinda Game Reserve). In Kruger National Park, 126km2 (three-male coalition) and 195km2 (single male) were recorded. Floater males have very large home ranges, averaging 777km2 (Serengeti National Park) to 1,390km2 (range 120–3938km2; single males, central Namibia) and 1,464km2 (range 555–4,348km2; coalitions, central Namibia). An Iranian male pair (likely floaters) used 1,737km2 in five months of radio-tracking. Cheetahs naturally occur at low densities, generally 0.3–3 per 100km2; estimates include 0.16 per 100km2 (Iran), 0.25−2 per 100km2 (Namibian farmlands), 0.5−2.30 per 100km2 (Kruger National Park, South Africa), 2 per 100km2 (Serengeti National Park, Tanzania) and 4.4 per 100km2 (Kgalagadi Transfrontier Park, South Africa). Temporary densities during migratory prey aggregations exceptionally reach 20 per 100km2 (Serengeti National Park, Tanzania).

Cheetahs are poorly built for arboreality but they easily scale large sloping trunks, sometimes as high as 5–6m. Cheetahs value trees as communal signposts for scent-marking, and for scanning the horizon for prey.

This wildebeest calf is no match for a Cheetah but its family is. Wildebeest and zebra herds often repel attacks on their juveniles by lone Cheetahs. They are much less likely to succeed in defending youngsters against Cheetah male pairs or trios.

Female Cheetahs with cubs spend much of their day scanning the environment. Vigilance by mothers is geared mainly towards finding prey except while feeding on kills, when females are keenly alert to the approach of other predators.

If caught, this Cheetah would be no match for the Lioness but adult Cheetahs easily outrun other carnivores provided they are not surprised. The risk of predation for them increases in dense woodland habitat where visibility is reduced.

Reproduction and demography

Cheetahs have a reputation for poor reproduction arising from the difficulties of breeding them in captivity, but wild cheetahs are prolific. Although Cheetahs have relatively low genetic variation, there is no evidence of reproductive inbreeding depression in wild Cheetahs. Cheetahs breed year-round, sometimes with weak birth peaks during prey lambing periods, for example November to May, Serengeti National Park. Iranian Cheetahs purportedly mate in winter (January to February) and give birth in spring (April to May), though there are few confirmed records. Oestrus is one to three days and mating is typically feline, though extremely rare to witness in the wild, often taking place in thick cover and at night. Gestation is 90−98 days. Litter size is typically three to six, rarely to eight; a litter of nine from Kenya may have included adoption. Inter-litter interval averages 20.1 months (Serengeti). Weaning begins at six to eight weeks and is complete by four to five months. Cubs reach independence at 12−20 months (average 17−18 months) and disperse as a sib-group. Females leave the group before sexual maturity while the males remain together. 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. Females can conceive at 21−24 months, first give birth around 24 months, averaging 29 months (Serengeti), and can reproduce up to 12 years. Males are sexually mature at 12 months, though rarely breed before three years.

Mortality 95 per cent of Serengeti cubs die before independence, most killed by Lions and less so Spotted Hyaenas, and most before eight weeks while cubs are still in the den. These are exceptionally high figures, and are likely restricted to the Serengeti’s very open plains and were recorded at a time when Lion densities were high. Cub mortality is lower everywhere else – for example, 37 per cent of Kgalagadi Transfrontier Park cubs survive to independence, and most mortality is due to predation (as for the Serengeti). Post-emergence (i.e. after the vulnerable denning period) survival estimates include 67 per cent (Kgalagadi Transfrontier Park), 62 per cent (Phinda Game Reserve), 57 per cent (Nairobi National Park, Kenya), 50 per cent (Kruger National Park) and 20 per cent (Serengeti). Infanticide by male Cheetahs has never been recorded, perhaps because the wide-ranging movements of females limit the benefits to males of killing unrelated cubs. Adult Cheetahs are occasionally killed by Lions, Leopards and Spotted Hyaenas, and territorial fights are a significant cause of mortality for males. Hunting accidents are rare but are sometimes fatal: a female pursuing Impalas in dense woodland was disembowelled while clearing a tree stump (Londolozi Game Reserve, South Africa) and a Cheetah hunting a male Grant’s Gazelle in the Serengeti died from injuries inflicted by the animal’s horns. Despite high genetic homogeneity, disease is rare in wild Cheetahs.

Lifespan Maximum 14 (average 6.2) years for Serengeti females; 11 (average 5.3) years for males; 21 in captivity.

Cheetah cubs play a game of ‘king-of-the-hill’. Baby Cheetahs belie their delicate appearance and, like all young cats, engage in very energetic and vigorous play.

Sport-hunting of Cheetahs and international trade of their skins is currently legal only in Namibia (where this photograph was taken) and Zimbabwe. There is also substantial illegal trade in live Cheetahs across borders, mainly in southern and north-east Africa.

STATUS AND THREATS

The Cheetah has disappeared from approximately 80 per cent of its historical range in Africa and its entire Asiatic range except for a single population of around 50 in central Iran. Total numbers are estimated at 7,000–9,300 adults and independent adolescents. Southern Africa (with roughly 4,500–5,000) and East Africa (with roughly 2,600) are the main strongholds; Namibia (~2,000), Botswana (~1,800) and Tanzania–Kenya (~1,700 combined) have the largest populations. Cheetahs are extinct or Critically Endangered in North and West Africa; fewer than 250 adults are estimated in two populations in Algeria and the protected WAP Parks Complex (Niger, Benin and Burkina Faso). They are otherwise thought to be extinct or relict across the Sahara and Sahel except for relict populations in southern Chad and southern Sudan.

Conversion of habitat to farmlands with replacement of prey by livestock is a key factor driving Cheetah declines. Across their range, a majority of Cheetahs occur outside protected areas where habitat conversion is prevalent and accelerating. Cheetahs are widely persecuted by livestock farmers despite causing relatively minor damage, and they are profoundly affected by loss of prey in pastoral areas. Human hunting of prey is a severe threat in the Sahel, North Africa and Iran where cheetahs are naturally very rare. Limited hunting for skins occurs, for example across the Sahel into southern Sudan where the market is mainly for luxury items such as traditional footwear called markoob. There is also significant illegal trade in live cubs and adults in north-eastern Africa, primarily from Somali ports to the Arab Gulf States where there is a high demand for Cheetahs as pets. Most cubs in this trade die in transit. High genetic homogeneity has had little impact on wild populations. The Cheetah is legally sport-hunted in Namibia (2013 quota: 150) and Zimbabwe (2013 quota: 50). CITES Appendix I, permitting trade of 205 hunting trophies and live animals. Red List: Vulnerable (global), Critically Endangered (Asia), Critically Endangered (hecki subspecies). Population trend: Decreasing.