Wild Cats of the World

IUCN RED LIST (2015):

Vulnerable (global)

Endangered (Asia)

Critically Endangered (West Africa)

Head-body length ♀ 158−192cm, ♂ 172−250cm

Tail 60−100cm

Weight ♀ 110−168kg, ♂ 150−272k

The Lion is one of the ‘big cats’ in the genus Panthera and is most closely related to the Leopard and somewhat more distantly to the Jaguar. The oldest Lion fossils are 3–3.5 million years old from East Africa.

More than 20 subspecies of Lions have been described in the past, based on largely superficial morphological differences and now considered invalid. Until recently, the most widely accepted taxonomy separated the Lion into Asian populations,

P. l. persica, and African populations, P. l. leo. However, genetic analyses strongly suggest the primary division between lion populations actually occurs in Africa roughly either side of the equator. An estimated 178,000 to 417,000 years ago, Lion range divided as repeated cycles of extremely wet and dry periods produced massive fluctuations in the extent of the Congo Basin rainforest and Sahara Desert. Neither habitat is suitable for Lions, which are thought to have retreated to two refugia, in West and Central Africa (or perhaps Asia) and in southern Africa. These two separate populations subsequently reoccupied suitable habitat during climatically stable periods when savannas expanded, coming together only relatively recently. The resulting dichotomy, reflected by strong genetic evidence, groups Asian populations with those in Central and West Africa (which would take the subspecies name P. l. leo, as the type specimen was described from North Africa within this range); and groups together populations from East and southern Africa (P. l. melanochaita). The same genetic analyses show evolutionarily significant subpopulations within the two main groups that are insufficiently different to be regarded as subspecies. In P. l. leo, there are three such units: Asia with the Middle East and North Africa (Lions are extinct in the latter two); West Africa west of the lower Niger River; and Central Africa. In P. l. melanochaita, there are three units: north-eastern Africa; south-western Africa; and East Africa with the rest of southern Africa.

The Lion is the largest carnivore in Africa and the second largest cat species. On average, Lions are comparable to Tigers in all measurements and, in fact, the Lion has a longer skull on average (around 1.2cm longer in females and 2cm longer in males).

The largest Tigers on record are slightly heavier and longer, while the smallest adult Tigers, from Sumatra, are considerably smaller than the smallest adult Lions. The Lion is a massive, robustly built cat with a deep chest, very powerful forequarters and heavily built limbs. The Lion is the most sexually dimorphic cat, both by secondary sex pelage (mane, belly fringe and elbow tufts in the male) and by size; male Lions are typically 30–50 per cent heavier than females. There is relatively little regional variation; Lions from India and the Sahelian savannas of West and Central Africa are 10–20 per cent smaller on average than Lions from southern and East African mesic savanna woodlands. Asiatic Lions usually have a distinct belly fold, which is only occasionally present in African Lions.

The Lion is uniformly coloured without body markings and is typically pale to dark tawny or sand coloured with cream or white underparts; lightening or deepening of the body colour produces variations including ash-grey, buff, light auburn and, rarely, dark brown. The backs of the ears are contrasting black with a scattering of silver hairs; the black ear-backs are distinct from a distance and may help Lions to locate hidden pride members spread out during a hunt. The tail ends in a distinct tassel of black or very dark brown fur, possibly useful as a flag for young cubs to follow in tall grass. White lions from the Kruger National Park region, South Africa, are leucistic (not albino) arising from a recessive gene for coat colour, and have pigmented eyes, nose and pads; they can be born to normally coloured parents. There are no records of melanistic lions. Lion cubs are born with dark brown rosettes, likely indicative of a more spotted, forest-dwelling ancestral species, which perhaps helps to camouflage young cubs in dens. The markings fade with age and are retained as faint spotting in some adults on the underparts, and, very rarely, covering the entire body (giving rise to historical accounts of new ‘spotted lion’ species).

The Lion is the only felid in which the males develop an extensive mane (that varies widely in colour and extent). Colour ranges from blonde to black, often with a corona of lighter colour surrounding the face. Mane growth begins at six to eight months, usually later in very hot climates. In mature males, the mane usually covers the entire head (excluding the face), neck, shoulders and upper chest; growth tends to be most extensive in mesic regions of southern and East Africa especially above 1,000m, sometimes extending along the ribcage and the belly. Captive individuals in northern hemisphere zoos with cold winters often grow extensive manes. The least developed manes occur in very hot climates, for example from Tsavo National Park, Kenya, to Niassa National Reserve, northern Mozambique (including Selous Game Reserve, Tanzania); Lions living at very low altitude here may be virtually maneless. Manelessness is also common in Sahelian populations. Male Asiatic lions have somewhat reduced manes, mainly around the face and crown. The mane is thought to signal the male’s relative genetic fitness to Lionesses and to intimidate rivals – mane length and colour convey information about male aggression and the ability to defend a pride from other males. Lionesses tend to choose males with the longest and darkest manes.

Similar species The Lion is one of the most recognisable mammal species and is quite distinct from all other cats. The unicolour Puma or Mountain Lion was named for its similar colouration to a Lioness, but the resemblance is otherwise weak and their ranges do not overlap.

The Lion has a patchy distribution in Africa south of the Sahara, chiefly in and around protected areas, and is restricted to a single Asian metapopulation in Gujarat state, India. The largest and most extensive Lion populations occur in East and southern Africa. They are severely reduced in most of Central Africa, with a fragmented distribution in northern Cameroon, southern Chad, Central African Republic, South Sudan and northern Democratic Republic of the Congo. Lions are extinct in most of West Africa where there are now only four populations in Senegal, Nigeria and a large tri-national population on the shared borders of Benin, Burkina Faso and Niger. The Lion is now extinct in North Africa, the Middle East and Asia except for the single population of around 400 in India.

Lions occur in a broad variety of habitats. They reach highest densities in mesic, open woodland and grassland savannas but they also occupy all kinds of moist and dry savanna woodlands, dry forest, scrub savanna, coastal scrub and semi-desert including very arid environments in the Kalahari and northern Namibia; they do not inhabit the interiors of true deserts and are absent from the Sahara. They traverse forest patches in savannas, such as the montane Harenna Forest in the Bale Mountains, Ethiopia, and Maramagambo Forest in Queen Elizabeth National Park, Uganda. Until very recently, they occurred widely in mosaics of equatorial forest and savanna, for example in Gabon and Republic of the Congo, but they are naturally absent from extensive moist, dense forest including the entire Congo Basin forest. The Indian population inhabits a mosaic of dry, deciduous teak forest and Acacia savanna. They occur from sea level to 3,500−3,600m (for example, on Mount Elgon and Mount Kenya, Kenya) and are recorded exceptionally moving to 4,200−4,300m (Bale Mountains, Ethiopia, and Mount Kilimanjaro, Tanzania). Except for largely unmodified livestock areas with low human densities and extant wild prey, Lions cannot occupy human-modified landscapes.

The Lion is a highly opportunistic and formidable predator. A single adult Lion is able to overpower much larger prey and, in prides, they are able to kill virtually everything they encounter; only healthy, mature male elephants are invulnerable to Lion predation. Lions are recorded eating insects to beached whale carcasses, but populations cannot persist without large herbivores weighing 60−550kg. In any given population, the diet is typically dominated by three to five ungulate species such as zebras, wildebeests, buffaloes,Giraffes, Gemsbok, Impalas, Nyalas, Greater Kudus, kobs, Thomson’s Gazelles, Chitals, Sambars and Warthogs. Small prey may dominate seasonally; for example, non-migratory Impalas and Warthogs are important prey to Lions during the lean, dry season in areas where large ungulates are migratory, including in Chobe National Park, Botswana, and Serengeti National Park, Tanzania.

Very large prey, including adult Common Hippopotamuses, rhinos and female elephants, are killed by large Lion prides usually when weakened through malnutrition, and a single lion is able to kill juveniles of these species. Over four years, 1993–1996, a pride in Chobe National Park, Botswana, consisting of two adult males, eight adult Lionesses and various cubs, killed 74 Savanna Elephants, including six adult females and one adult bull (previously injured by another bull). As elephant densities increase, for example in Chobe and Hwange National Parks, the proportion of elephant calves in Lion diet increases. Besides large ungulates, practically any species may be eaten by the Lion but none contributes significantly to the diet. Lions are recorded killing Aardvarks, porcupines, primates, including gorillas, Chimpanzees and baboons, many bird species including Ostriches, reptiles including large Nile Crocodiles and African Rock Pythons, fish and a wide variety of invertebrates. Lions regularly kill other carnivore species including Leopards, Cheetahs, hyaenas, African Wild Dogs, Cape Fur Seals and a wide variety of smaller species. Carnivores are apparently unpalatable to Lions and are generally left uneaten when killed. Cannibalism occasionally occurs, most often of cubs killed by infanticidal male Lions. Although Lions go for long periods killing only wild prey even when domestic livestock is abundant in an area, they do prey on livestock including cattle, goats, sheep, donkeys, horses, camels and Water Buffaloes (India), and are occasionally recorded killing domestic dogs. Lions rarely eat people although isolated, localised pockets of persistent man-eating occur very rarely, for example in areas of south-eastern Tanzania and northern Mozambique, where Lions locally regard people as prey. An estimated 50–100 people are killed annually by Lions in remote communities in these areas.

Hunting is mainly nocturno-crepuscular and terrestrial. Lions are too heavy to climb well, and do not hunt arboreally, although prey (for example, baboons and guineafowl) is sometimes snatched from lower branches. Lions move and hunt communally, often with the entire pride in attendance, including cubs older than four to five months. Young cubs are left behind, sometimes in the company of older cubs or males. Adult females typically initiate most foraging bouts and they make the most kills. However, male Lions are capable hunters that regularly make their own kills when unaccompanied by the females and males increase the success rates of prides hunting very large prey, for example buffaloes, Giraffes and elephants. Lions search for prey as they walk, using their excellent vision and hearing to identify potential targets; or they rest around profitable ambush sites such as waterholes until a suitable opportunity arises. Most hunts begin with a careful stalk by one or more members of the pride to as close as 15m from the prey, followed by an explosive rush. The Lion’s top speed is an estimated 58kmph, which it can maintain only for around 250m, and most chases are shorter. The exception occurs during hunts of buffalo herds that often stand their ground. Lions harass the herd until it stampedes, providing opportunities for the Lions to isolate a straggler during the ensuing chase, which often covers up to 3km with an exceptional record of 11km. Lions kill large prey by a suffocating throat bite administered by an adult, often while the rest of the pride begins feeding.

Lions hunt communally but the degree of actual cooperation is exaggerated. On occasion, a single Lioness stalks prey while the rest of the pride watches, joining her only when the prey is caught. More often, multiple females are involved in which they fan out around the prey and one or more individuals gives chase; the capture is often the result of fleeing prey running into a Lioness lying in wait. During hunts of very fast Springbok in open habitat in Etosha National Park, Namibia, Lionesses apparently assume a specific position and coordinate their movements, with ‘wing’ positions driving Springbok to ‘centre’ positions. Cooperative hunts are more successful than hunts by lone Lionesses, and hunts are more successful when each Lioness occupies her preferred position. Cooperative hunting is also more prevalent during hunts of large dangerous prey, especially buffaloes and elephants. Estimates of hunting success include: 15 per cent (Etosha National Park, Namibia), 23 per cent (Serengeti National Park, Tanzania) and 38.5 per cent (Kalahari, South Africa). Lions readily scavenge, and this comprises 5.5 per cent of intake in Etosha National Park to almost 40 per cent in Serengeti National Park. They frequently appropriate kills from other carnivores.

The Lion is highly social and is the only felid that forms large, mix-sexed prides comprising multiple generations. Lions are almost constantly in the company of other pride members and rarely alone except when females temporarily abandon the pride to give birth. The basis of the pride is a matriline of 1−20 (usually three to six) related Lionesses that communally defend a territory and raise their cubs. Each pride usually has a coalition of one to nine (usually two to four) adult males that typically immigrate from other prides and are unrelated to the breeding females. Pride size is only weakly correlated to prey abundance (whereas this strongly influences home range size and Lion density) and is remarkably consistent over much of the East and southern African range – typically three to six females and two to three males. This would probably also apply in West and Central Africa if it wasn’t for very high levels of human persecution of Lions and prey. Pride size exceptionally and temporarily reaches 45−50 including cubs in optimal conditions but invariably decreases as large cohorts of subadults disperse.

Female membership of the pride is stable but small subgroups often come and go within the pride range in a ‘fission-fusion’ pattern so that the entire pride is rarely all together. Females usually stay with the pride for life. They occasionally disperse following a pride takeover by immigrant males, or to avoid mating with male relatives. Young males are evicted or leave the pride at 20−48 months, entering a protracted transient phase lasting up to three years before attempting to acquire their own pride. Coalition males are usually related to each other but male singletons and pairs often join with unrelated males during dispersal to form a coalition. Whether related or not, coalition members have close bonds and remain together for life, defending their territory and females from male intruders. Immigrant males challenge resident males for access to female prides, sometimes resulting in violent fights that are often fatal. Lionesses participate in pride defence and large female groups may succeed in repelling incoming coalitions, though Lionesses are occasionally killed by strange males in these encounters. Inter-pride territorial skirmishes are also common, sometimes resulting in neighbouring Lionesses killing each other or cubs. When new males succeed in taking over a pride, they usually kill or evict all unrelated cubs younger than 12−18 months to hasten the Lionesses’ return to oestrus. Depending on coalition strength, males may take over a neighbouring pride while still defending their current pride. Coalition tenure is generally two to four years.

Territories are very stable, with female matrilines remaining in essentially the same area for many generations. Territory size varies widely, depending on the productivity of the habitat and therefore the available prey biomass. Pride territories in Gir Protected Area (India) where there is a superabundance of Chital deer are very small at 12−60km². Serengeti (Tanzania) pride territories average 65km² (woodlands) to 184km² (grasslands), reaching a maximum of 500km². Average range size in West and Central Africa (based on very few prides) ranges from 256 km² in relatively well-watered Pendjari National Park, Benin, to 756km² in drier Waza National Park, Cameroon. In semi-arid savanna in Hwange National Park, Zimbabwe, female prides have an average range size of 388km²(range, 35−981km²) and slightly more for male coalitions, averaging 478km²(71−1002km²). Range size is very large in arid areas; for example, 1,055−1,745km²(Khaudom Game Reserve, Namibia), 266–4,532km² (Kgalagadi Transfrontier Park, South Africa) and 2,721−6,542km² (Kunene, north-western Namibia). Two Kunene male coalitions (possibly nomads) had ranges of 13,365–17,221km². Density estimates include 0.05−0.62 per 100km² (Kunene, Namibia), 1.5–2.0 per 100km² (Kalahari, South Africa), 3.5 per 100km² (Hwange National Park, Zimbabwe), 6−12 per 100km² (Kruger National Park, South Africa ), 12−14 per 100km² (Gir Protected Area, India) and up to 38 per 100km² (Lake Manyara National Park, Tanzania).

The Lion breeds aseasonally, though births often peak when seasonally breeding ungulates give birth, for example March–July in East Africa (Serengeti National Park, Tanzania) and February–April (Kruger National Park, South Africa). Oestrus averages four to five days and gestation is 98−115 days, averaging 110 days. Litter size is typically two to four cubs, exceptionally up to seven. Lionesses usually leave the pride to give birth, and cubs are kept isolated until around six to eight weeks old, perhaps to insulate them from rough play or accidental deaths by other pride members. Females in the same pride often give birth synchronously and communally care for cubs; females suckle all cubs but they only transport their own by holding them gently in the mouth. Weaning begins around six to eight weeks but suckling may continue to eight months. Inter-litter interval is typically about three years in savanna-woodland ecosystems with resident prey, and declines to 20−24 months in areas with migratory prey, where cub survival rates are consequently lower. When reintroduced into areas from which they have been previously extirpated, Lions breed at similarly high tempos to migratory systems. Cubs can hunt independently at around 18 months but they rarely disperse before two years, and typically leave the pride at just under 36 months of age when their mothers initiate the next breeding event (in most savanna-woodland habitats).

In migratory prey systems and arid habitat, dispersers wander extensively, often ending up more than 200km from their natal range. However, in wooded savannas with abundant resident prey (especially buffalo), most young males remain within or close to their natal range for several years and may establish territories close to their original pride. Males are more likely to survive dispersal if they leave the pride as old as possible. All dispersers younger than 31 months monitored between 1999 and 2012 in Hwange National Park, Zimbabwe, did not survive. Of 49 dispersers whose fates were known, 65 per cent (17 males and 15 females) survived to establish a territory, while 26 per cent were killed by people. A male Lion in this population that dispersed at 27 months was transient for 848 days and travelled 4,223km before he was shot for killing livestock. Lionesses can conceive at 30−36 months but typically first give birth around 42−48 months and cease reproducing after age 15. Males are sexually mature at 26−28 months but rarely breed before five to six years of age. In areas where adult males are depleted (typically due to excessive trophy hunting), very young males of two to four years replace these older mature males as the main breeding males in the population.

Mortality Cub mortality varies widely, depending on the region and seasonal or annual fluctuations in prey availability. In the first year, around 16 per cent of cubs die in southern Kruger National Park, South Africa, where there is abundant, resident prey. In the arid Kalahari, this figure is around 40 per cent. It is highest in areas with dramatically fluctuating prey levels; for example, with migratory systems such as in Serengeti National Park, Tanzania, where an average of 63 per cent of cubs die in the first year. Cubs die mostly from infanticide (which is higher in the Serengeti than anywhere else), predation (mainly by Leopards and Spotted Hyaenas) and starvation due to food shortages. Annual cub mortality in the second year drops dramatically; for example, to 20 per cent (Serengeti) and 10 per cent (Kruger). Male Lions experience much higher rates of mortality during dispersal, resulting in a typical adult sex ratio of two to three Lionesses per adult male. Apart from human-caused mortality, adult Lions die mainly in fights with other Lions (especially males), from injuries sustained while hunting large prey (especially buffaloes) and from starvation when old or debilitated. Lions often survive catastrophic injury from fights or hunting accidents, usually to the lower spine and hindlegs. These Lions seldom live to an old age. Disease is uncommon but the intense sociality of Lions fosters disease transmission, and episodes are occasionally severe; more than 1,000 lions (40 per cent of the population) died during a 1993−4 canine distemper outbreak in the Serengeti.

Lifespan 18 years for females, 16 (but rarely over 12) for males in the wild and 27 in captivity.

STATUS AND THREATS

The Lion reaches high densities in well-protected, productive habitat, but it has undergone a massive range collapse and continues to decline across most of its remaining distribution. The species is extinct in Asia except in Gujarat, western India, where about 300 live in the Gir Protected Area (1,883km2) and another approximately 100 live in seven small satellite populations nearby. In Africa, the most optimistic estimate of Lion distribution is 16.3 per cent of its historical range, counting poorly known areas where its continued presence is uncertain but possible. The Lion is unequivocally known from slightly less than 8 per cent of its original range. It is extinct in 15 African countries and possibly extinct in an additional seven. The latest estimate of approximately 32,000 Lions is overly optimistic; the number is probably closer to 20,000, many of which occur in small, isolated and declining populations. There are only eight countries thought to contain at least 500 adults: Botswana, Kenya, Mozambique, Namibia (possibly), South Africa, Tanzania, Zambia and Zimbabwe. The Lion is Critically Endangered in West Africa where the number of adults is thought to be fewer than 250.

The Lion continues to decline over most of its range. Overall, the total population is estimated to have declined 38 per cent since 1993 but this conceals a far more severe decline across most of the range. Five countries (Botswana, India, Namibia, South Africa and Zimbabwe) have stable (or nearly stable) and increasing populations where the total population is estimated to have increased 25 per cent since 1993. Such increases in a relatively small part of the range disguise the severity of the decline everywhere else, which is estimated at 59 per cent since 1993. Accordingly, although the Lion is classified as Vulnerable, it qualifies to be considered Endangered in most of its range.

Lions have declined mainly from the extensive loss of habitat and prey to agriculture and livestock herding, coupled with pervasive eradication by people. They are persecuted intensely by herders and suffer from widespread official killing of ‘problem’ animals. Their habit of scavenging makes them highly vulnerable to poisoned baits and to snares set for bushmeat. Sport-hunting is legal in 13 African countries, taking approximately 600−700 (mainly males) annually, and contributes to population declines when poorly managed. Genetic impoverishment of small, isolated populations possibly leads to declines and vulnerability to disease.

CITES Appendix II. Red List: Vulnerable (global), Endangered (Asia), Critically Endangered (West Africa). Population trend: Decreasing.