Lion

Name and taxonomy

Carl Linnaeus named the lion Felis leo in 1758. Lorenz Oken created the genus name Panthera in 1816, and the lion moved into it later. The English word lion reached the language through Anglo-Norman from the Latin leo, which itself came from Greek. iNaturalist calls the genus Panthera the roaring cats, and the lion shares it with the tiger, leopard, jaguar and snow leopard.

Between the mid 1700s and the mid 1900s, naturalists proposed 26 lion subspecies. Most were separated only by the size and colour of the mane and coat. In 2017 the Cat Specialist Group cut the list to two subspecies. P. l. leo covers the Asiatic lion, the extinct Barbary lion, and the lions of West and northern Central Africa. P. l. melanochaita covers the lions of East and Southern Africa, together with the extinct Cape lion. Genetic studies split these two groups into seven smaller clades.

Evolution and relatives

The Panthera line split from the rest of the cat family about 10.8 million years ago. The oldest fossils that are clearly lions come from Olduvai Gorge in Tanzania and may be 2 million years old. Lions spread out of Africa into Eurasia about 1 million years ago. That wave gave rise to the cave lion, Panthera spelaea, which is now treated as a separate species. The cave lion lived across Europe, Siberia and into Alaska, and it died out by 11,900 years ago. Its American cousin, Panthera atrox, left the cave lion line about 165,000 years ago and ranged from Canada far to the south.

Expanding rainforest cut the East and Southern lions off from the West and North lions between 183,500 and 81,800 years ago. Lions from North Africa moved into southern Europe and on into Asia, starting about 38,800 years ago and ending near 8,300 years ago. In zoos, lions have bred with tigers to produce ligers and tigons, and with leopards to produce leopons.

Appearance

The lion is a heavy, deep-chested cat with a short round head, round ears and a tufted tail. Males weigh about 189 kilograms on average, and the heaviest on record reached 272 kg. A female averages about 126 kg. Head, body and tail together measure 240 to 330 centimetres, and the tail alone is 0.6 to 1.0 metres. Skeletal muscle makes up 58.8% of body weight, the highest share known in any mammal.

The coat runs from pale buff through silvery grey and yellowish red to dark brown, with paler underparts. Cubs are born with dark spots on a greyish coat, and faint spots can persist on the belly and legs of adults. White lions turn up around Kruger National Park in South Africa. They carry a recessive gene for leucism, so their eyes and skin keep normal pigment, unlike an albino.

The male's mane sets the lion apart from every other cat. It starts to grow when testosterone rises in adolescence and reaches full size at about four years. Darker manes go with higher testosterone, longer tenure in a pride and more cubs, but also with more heat stress. The mane probably evolved to signal male condition to females rather than to protect the neck in fights.

Habitat and range

Lions favour open country: grassy plains, savanna, thorn scrub along rivers and light woodland. They avoid dense closed forest and true desert. Savanna with 300 to 1,500 millimetres of rain a year makes up most of their African habitat. Lions climb high: they are recorded at 3,600 metres on Mount Elgon and at 4,240 metres in Ethiopia's Bale Mountains.

Historically the lion ranged across all of Africa outside the Congo rainforest and the Sahara. Today the species lives in fragmented populations, mostly inside protected areas in East and Southern Africa. West African lions are cut off from those in Central Africa and number only about 400. GBIF holds records of the species from 71 countries, but that count includes zoos and escapes, not the native range.

In Asia the lion once ranged from Greece through the Caucasus and Iran to central India. The only Asiatic population left lives in Gujarat, in Gir National Park and the dry scrub forest around it.

Diet and hunting

The lion is a hypercarnivore that takes a wide range of prey, and it sits at the top of its food web. Its main prey are medium and large hooved animals: zebra, wildebeest, buffalo, gemsbok, giraffe and warthog. In India the usual wild prey are chital and sambar deer, while livestock makes up a large share of kills outside the park. Lions also scavenge, and they watch for circling vultures as a sign of a carcass.

Females do most of the hunting, because a maned male is easier for prey to spot. In the Serengeti a lone hunter succeeds on 17% of attempts, and a group on 30%. Lions lack stamina, so they stalk to within about 20 metres before the rush. The charge reaches 48 to 59 kilometres per hour, and one lion was clocked at 74.1 km/h in 2018. The kill comes from a bite that clamps the throat or muzzle until the animal stops breathing, up to 13 minutes.

A lioness needs 5 kilograms of meat a day on average, and a male needs about 7 kg. After a big kill a lion can eat up to 30 kg in one sitting, then rest for hours before it returns. Males squabble most at a carcass, and cubs go hungry first when food is short.

Behaviour and social life

A pride holds on average about 15 lions: several adult females, up to four adult males, plus cubs. Females stay in the pride where they were born, so pride females are close relatives. Members split into small parties and rejoin, and they keep contact by roaring. Female pride mates do not fight over food or rank, which makes shared cub care possible.

Young males leave their birth pride at about two to three years of age. They wander as nomads, often with a brother, and form a coalition to take over a pride. In the Serengeti, a coalition first wins a pride when its males are 3.5 to 7.3 years old. A coalition of two rarely holds a pride longer than two and a half years. The pride males patrol the edges of the territory and take the lead against intruders.

Lions rest for about twenty hours a day. Activity picks up after dusk with grooming and socialising, and hunting peaks late at night and towards dawn. Where prey is dense, open country supports up to 12 lions per 100 square kilometres.

Communication and senses

The roar is the lion's long-distance signal, and it carries about 8 kilometres on a still night. A male's roar is louder and deeper than a female's. Pride members often roar in chorus, which may strengthen the bond between them. Other calls include growls, snarls, puffs, hums and a breathy purr, and cubs mew and bleat.

Close up, lions greet by rubbing heads and licking each other's head and neck. Males spray urine on bushes and scrape the ground to mark the territory. When a lion sniffs a scent mark it pulls a grimace, with open mouth and wrinkled nose, called the flehmen response. Like other cats, the lion sees well in low light, which suits its night hunting.

Reproduction and life cycle

Lions breed in any month, though births in Africa peak in the rainy season. A female is in heat for four days and mates on average 2.2 times an hour during that time. Gestation lasts about 110 days. The mother gives birth away from the pride in a thicket, reed bed or cave. A litter usually holds two to four cubs, sometimes as many as six, each 1.2 to 2.1 kilograms at birth.

Cubs are born blind, open their eyes after about a week and walk at three weeks. She brings them to the pride at six to eight weeks. Pride females often give birth close together, and every cub can suckle from any nursing female. Weaning ends at six to seven months, but cubs depend on the pride until at least 16 months.

A new male coalition usually kills the unweaned cubs of the previous males. The mothers then return to heat within two to three weeks, which lets the new males father cubs sooner. Females first breed at about four years and males at five, though males peak at five to nine. Wild lions live 12 to 17 years, and females usually outlive males. The record in captivity is 30 years.

Predators and threats

An adult lion has no natural predator. Most adults die violently all the same, killed by other lions or by people. Spotted hyenas take cubs, and they also kill sick or old adults. Nile crocodiles sometimes kill a lion at the water, and lion claws have been found inside crocodiles. Infanticide by incoming males remains the biggest single killer of cubs.

Ticks, tapeworms and biting flies trouble lions, and a 1962 stable fly outbreak cut the Ngorongoro population from 70 to 15. Beyond disease, the real threats are the loss of habitat and prey and conflict with herders over livestock. Snares, poisoned carcasses and rifles kill lions that leave protected land. Trophy hunting is legal in six African countries, and the killing of Cecil in 2015 made the practice a public argument.

Population and conservation

The IUCN Red List rates the lion as Vulnerable. The listing dates from 1996, after surveys showed African populations fell by about 43% from the early 1990s. Current estimates put the wild population near 20,000. Estimates for Africa in 2002 to 2004 ran from 16,500 to 47,000, which shows how uncertain the counts are.

The West African population is listed as Critically Endangered, with about 400 animals and fewer than 250 adults. Most of them live in the W-Arly-Pendjari complex, where Benin, Burkina Faso and Niger meet. In 2015, two lions appeared in Ghana's Mole National Park, the first there in 39 years. The same year a camera trap found up to 200 lions in Alatash, Ethiopia, a population thought lost.

South Africa rates its own lions as Least Concern, because they live in well-managed reserves. The Asiatic lion is listed as Endangered on its own. Its numbers rose from 180 in 1974 to 400 in 2010, then 523 in 2015 and about 650 in 2017. Lion Conservation Strategies drawn up in 2005 aim to keep habitat, protect wild prey and reduce conflict with people. Where herders keep cattle in strong enclosures at night, lions kill far fewer livestock.

Relationship with humans

People have kept lions since Roman times, and zoos have shown them since the late 18th century. Zoos and wildlife parks hold over 1,000 African lions and around 100 Asiatic ones. In 1987 most zoo lions in North America turned out to be African-Asiatic hybrids, so breeding was reset with wild-born imports. Lion taming began in circuses in the early 19th century with Henri Martin and Isaac Van Amburgh.

Lions rarely hunt people, and a lion that meets a person on foot usually moves off. Some do turn to man-eating, most of them males. In 1898 two Tsavo lions took 28 recorded railway workers in nine months in Kenya. Attacks in rural Tanzania rose sharply from 1990 to 2005, with at least 563 people attacked. Worn teeth explain some cases, but a shortage of wild prey near villages is the more likely cause.

Kings hunted lions for show in Egypt, Assyria and Mughal India. Maasai men once killed a lion as a rite of passage, but elders now discourage solo hunts. Today the lion is one of the most valuable species for ecotourism in Africa. Trophy hunting continues in six countries, and white lions have been bred for canned hunts in South Africa.

In culture and myth

The lion is called the king of beasts in many languages, and it stands for power, royalty and protection. Rulers have taken its name, from Sundiata Keita, the Lion of Mali, to Richard the Lionheart. In Egypt the lion belonged to the sun and the Nile, and the war goddess Sekhmet had a lioness head. Nubia worshipped the lion god Apedemak, tied to the flooding of the Nile. In Mesopotamia the lion was the beast of the goddess Inanna, and glazed lions line the Processional Way to the Ishtar Gate.

In the Bible the lion marks the tribe of Judah, and Daniel survives a night in a lions' den. Ethiopian emperors took the Lion of Judah as the sign of the Solomonic line. Hindu tradition tells of Narasimha, the man-lion form of Vishnu, who kills the demon king Hiranyakashipu. Emperor Ashoka's lion capital, with four lions back to back, dates from the 3rd century BCE. China never had wild lions, yet stone lions guard palace and temple gates and the lion dance is over a thousand years old.

The Greeks told of the Nemean lion, whose hide no weapon could pierce, so Heracles strangled it and wore the skin. Aesop's fable of the lion and the mouse shows a small favour repaid. African folklore often makes the lion strong but slow-witted, and a smaller animal tricks it. Some West and East African traditions make the lion the link between seers and the spirit world, and a healer's animal. Modern stories run from the Cowardly Lion in The Wonderful Wizard of Oz to Aslan and The Lion King.