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Lifespan, longevity, and ageing

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Maximum longevity: 27 years (captivity) Observations: Some older publications list lions as capable of living up to 30 years (Theodor Haltenorth and Helmut Diller 1977), which is based on anecdotal reports and probably incorrect. Record longevity in captivity belongs to a wild born male that died at Cologne Zoo at about 27 years of age. A hybrid between a leopard and a lion lived for 24 years and a hybrid between a lion and tiger lived for 24.2 years (Richard Weigl 2005).
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Associations

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Adult lions have no natural predators, excepting persecution by humans. Lions often kill and/or compete with other predators (leopards Panthera pardus and cheetahs Acinonyx jubatus). Spotted hyenas Crocuta crocuta defend kills or scavenged food from immature and female lions, but typically leave the food to a big male lion. Hyenas are known to kill lion cubs, juveniles, or weak and sick adult lions.

Lion cubs, if left alone, can be vulnerable to other large predators. However, infanticide is the primary threat to cubs.

Human poaching is a problem for lions. These animals are poached with wire snares, rifles, and arrows. Since lions are scavengers, they are particularly vulnerable to intentionally poisoned carcasses. There are still poachers that operate within some national parks in Africa. It has been estimated that in the 1960's, poachers were responsible for approximately 20,000 lion deaths per year in Serengeti National Park. Trophy hunting is allowed in 6 African countries.

Known Predators:

  • spotted hyenas (Crocuta crocuta)
  • humans (Homo sapiens)
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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Morphology

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Lions are large cats with short, tawny coats, white underparts, and long tails with a black tuft at the end. They are sexually dimorphic and male lions are the only cats with manes. Three year-old male lions grow manes that vary in color from black to blond. Manes tend to be fuller in open habitats. Adult males typically weigh 189 kg; the heaviest male on record weighed 272 kg (Mount Kenya). Females weigh 126 kg on average. The average male height is 1.2 m and the average female height is 1.1 m. Length ranges from 2.4 to 3.3 m and tail length ranges from 0.6 to 1.0 m; the longest male lion recorded was 3.3 m.

Cubs have brown spots on a grayish coat until the age of three months; spots may remain on stomach, especially in east Africa. Albinism does occur in some populations, but there are no published records of melanism (black fur) in lions. Adult lions have 30 total teeth and adult females have four mammae.

Asiatic lions (P. l. persica) are slightly smaller than African lions and have shorter manes, thicker elbow and tail tufts, and longitudinal skin folds on their stomachs. Although Asiatic lions are genetically distinct from African lions, the genetic difference between the two species is smaller than that between human races.

Range mass: 126 to 272 kg.

Range length: 2.4 to 3.3 m.

Sexual Dimorphism: male larger; ornamentation

Other Physical Features: endothermic ; bilateral symmetry

Average basal metabolic rate: 94.58 W.

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Life Expectancy

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Female lions typically live longer than males. Males reach their prime between five and nine years but few males survive past ten years of age. Some males have survived until 16 in the wild. Females normally live until 15 or 16 years. In the Serengeti, females live up to 18 years. In captivity, lions live approximately 13 years. The oldest known lion was 30 years old.

Adult lions have no predators, but are vulnerable to humans, starvation, and attacks from other lions. Infanticide is an important contributor to cub mortality and cub mortality increases when prey is scarce.

Female Asiatic lions live an average of 17 to 18 years, with a maximum of 21 years. Male Asiatic lions generally live for 16 years. Adult Asiatic lions have a less than 10% mortality rate. In the Gir forest, 33% of cubs die during their first year of life.

Range lifespan
Status: wild:
18 (high) years.

Range lifespan
Status: captivity:
30 (high) years.

Average lifespan
Status: wild:
14 years.

Typical lifespan
Status: captivity:
30 (high) years.

Average lifespan
Status: captivity:
13 years.

Average lifespan
Status: wild:
15.0 years.

Average lifespan
Status: wild:
29.0 years.

Average lifespan
Status: wild:
15.0 years.

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Habitat

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African lions live in plains or savanna habitat with a large prey base (mostly ungulates) and sufficient cover available. In these optimal habitats, lions are the second most abundant large predator, after spotted hyenas Crocuta crocuta. Lions can also live, with wider ranges, in most habitats except in tropical rainforests and in deserts. Lions can live in forested, shrubby, mountainous, and semi-desert habitats. They are capable of living at high altitudes. There is a lion population in the Bale Mountains of Ethiopia at 4,240 m.

Asiatic lions live in scrubland and teak forest in the small Gir Forest preserve of India.

Range elevation: 4240 (high) m.

Habitat Regions: temperate ; tropical ; terrestrial

Terrestrial Biomes: savanna or grassland ; forest ; scrub forest ; mountains

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Distribution

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African lions (Panthera leo) live in most of sub-Saharan Africa except in desert and rainforest habitats. Lions were once exterminated from South Africa, where they remain in Kruger and Kalahari Gemsbok National Parks and possibly some other protected areas. Lions once ranged throughout southwest Asia and north Africa. Asiatic lions (P. l. persica) belong to the single remaining subspecies in this region. Once roaming from Greece to central India, Asiatic lions persist in the Gir forest of northwest India.

In addition to the asiatic subspecies, many taxonomists contend that there are five extant African subspecies. Each subspecies is identified by geographic region. Panthera leo senegalensis (west African or Senegalese lions), P. l. azandica (north east Congo lions), P. l. bleyenberghi (Katanga, Angolan, or south Congo lions), and P. l. krugeri (south African or Transvaal lions). Panthera leo krugeri includes Kalahari lions (sometimes denoted as P. l. verneyi). Lastly, there are East African lions (P. l. nubica). These animals have been categorized as Somali lions (P. l. somaliensis), Masai lions (P. l. massaicus), Serengeti lions (P. l. massaicus), Congo lions (P. l. hollisteri), and Abyssinian lions (P. l. roosevelti). It should be noted, however, that there is some debate as to the validity of the African subspecies classifications, leaving only the Asiatic subspecies, P. l. persica, uncontested.

Biogeographic Regions: palearctic (Native ); oriental (Native ); ethiopian (Native )

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Trophic Strategy

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Lions are predatory carnivores. They usually hunt in groups, but the actual killing is done by an individual lion. They frequently bring down prey much bigger than they are themselves. Showy males have more difficulty hunting than females because of their conspicuousness, therefore females in a pride do the majority of hunting. Males are still more aggressive during feeding than are females, even though they are less likely to have killed the prey.

African lions eat the most common large ungulates in the area (Thompson's gazelles Eudorcas thomsonii, zebras Equus burchellii, impalas Aepyceros melampus, and wildebeests Connochaetes taurinus). Individual prides tend to have their own eating preferences. Some prides tend to target large prey such as cape buffalo Syncerus caffer and giraffe Giraffa camelopardalis. Lions that are not able to capture large prey will eat birds, rodents, fish, ostrich eggs, amphibians and reptiles. Lions also actively scavenge, taking cues from hyenas and vultures.

In Tanzania's Serengeti National Park, local lions subsist on a diet comprized mainly of 7 species: zebras Equus burchellii, wildebeests Connochaetes taurinus, Thompson's gazelles Eudorcas thomsonii, buffalos Syncerus caffer, warthogs Phacochoerus aethiopicus, hartebeests Alcelaphus buselaphus, and topis Damaliscus lunatus provide 90% of their diet.

Hunting effectiveness is increased by hunting in groups. Serengeti research has shown that individual lions succeed in their hunting 17% of the time, whereas group hunts succeed 30% of the time.

Animal Foods: birds; mammals; amphibians; reptiles; fish; eggs; carrion

Primary Diet: carnivore (Eats terrestrial vertebrates)

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Associations

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Lions are the top predators in their range. It is not clear to what extent lions regulate their prey population; some studies have shown that food availability plays a larger role in regulating prey populations than consumption by lions.

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Benefits

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Lions are a glamorous species, well-known throughout the world. They are a cultural icon in England and are one of the highest valued eco-tourism species in Africa. They are also the subject of many documentaries and research efforts.

Positive Impacts: ecotourism ; research and education

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Benefits

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Humans are fearful of being attacked by lions and of their livestock being killed by lions. In most cases, this has not been a large problem. Lions have historically coexisted with the Masai and their cows in east Africa. When the prey base is plentiful, lions do not typically attack cattle. Also, if a lion sees a human on foot, it will typically run in the opposite direction.

There have been well-publicized attacks on humans by lions, such as the man-eating lions of Tsavo (killed 135 construction workers) portrayed in the 1996 movie The Ghost and the Darkness. As lions lose habitat, they are more frequently entering human inhabited areas, thus creating more conflict and potential for "problem lions" that attack humans.

Lions are frequently infected with FIV (Feline Immunodeficiency Virus) which is similar to HIV. In Tanzania’s Serengeti National Park, Ngorongoro Crater, and Kruger National Park of South Africa, 92% of lions tested had FIV. FIV does not seem to adversely affect lions but it will kill domestic cats.

Negative Impacts: injures humans (bites or stings); causes or carries domestic animal disease

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Conservation Status

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Panthera leo leo (Barbary lions) and P. l. melanochaita (Cape lions) are two extinct subspecies of African lion.

African lion populations have greatly declined in West Africa and in many African countries they are restricted to protected areas. If there are no connecting corridors between wildlife reserves genetic viability will likely become a problem.

Asiatic lions, P. l. persica, are confined to one population in the Gir Forest reserve of India, consisting of less than 200 mature individuals. This subspecies is listed as critically endangered on the IUCN red list and is on Appendix I of CITES. Another population of Asiatic lions is desperately needed in order to safeguard the survival of this subspecies. The Palpur-Kuno Wildlife Sanctuary in northern Madhya Pradesh had been identified as a potential reintroduction site in India. Threats to the Gir Forest population include the close proximity of humans and their cattle and habitat degradation.

Some very small lion populations require genetic management in order to survive and maintain genetic diversity. Hluhluwe-Umfolozi Park (HUP) in Natal, for example, has a population of 120 lions produced from only three lions that were introduced into the park in the 1960s. In 2001, researchers tried artificial insemination techniques to rejuvenate the genetic pool of these South African lions. This process is very difficult and energy intensive. Inbred populations could also potentially be rejuvenated by introducing adult females and whole prides into an area (minimizing conflict between existing lions and introduced lions).

US Federal List: no special status

CITES: appendix i; appendix ii

IUCN Red List of Threatened Species: vulnerable

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Behavior

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Lions have the cognitive ability to recognize individuals and to interact with other lions to benefit their own survival. They use visual cues in this communication. For example, the mane is thought to signal to other lions the sex of a male from a distance and to indicate individual fitness. (The rate of mane development is mostly controlled by testosterone.)

Resident male lions regularly mark their territory by spraying vegetation with urine and by scuff-marking. Females spray occasionally. This behavior starts at the age of 2 years. This type of marking uses both chemical and visual communication signals.

Male lions start to roar at 1 year of age and females start shortly after this. The male’s roar is both louder and deeper than the female's. Lions can roar at any time, but they typically stand or crouch while roaring. Roaring serves to advertise territories, to communicate with other pride members, and to demonstrate aggressions toward enemy lions. Lions also roar in chorus; this may be a form of social bonding. This is accoustic communication.

Finally, lions use tactile communication. Males engage in physical aggression during pride take over. There is touching during greetings between pride members. Physical communication passes between lactating females and the cubs they are nursing.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: choruses ; scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Untitled

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Fortunately, individual lions can be dependably identified in an unobtrusive manner. Lion whisker spots are similar to our finger prints. Every individual lion has a unique whisker spot pattern. More specifically, the number and relative position of whisker spots on the top row are used to identify individuals.

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Reproduction

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Lions breed year-round and are usually polygynous. It is estimated that lions copulate 3,000 times for every cub that survives over one year. One estrus out of every 5 results in a litter and lions mate approximately 2.2 times per hour for the 4 day estrus period. The first male member of a pride that reaches a female in heat has the mating priority over her. Fighting between pride members over females does not normally occur.

Male lions are conspicuously large and showy because they have the opportunity to control the reproduction of many females when they rule over a pride. Males form coalitions with each other to increase their chances of pride takeover. The fierce competition among males and the social structure of a pride have led to infanticide by both males and females. Successful males that takeover a pride have about 2 years before another younger, stronger coalition will replace them. Pride takeover battles are often violent leading to severe injury or death of the losing lions.

It is to the successful male’s reproductive advantage to kill the suckling cubs of the defeated males. A nursing lioness that loses her cubs will come back into estrous within 2 to 3 weeks. The normal time between births is 2 years, which is the typical time for a male to rule a pride. Therfore, by killing all unweaned cubs at the time of pride takeover, males can ensure that they have some opportunity to father offspring of females who would otherwise not be available to them during their tenure as pride leaders. Females vigorously defend their cubs during a takeover and are sometimes killed also.

Mating System: polygynous ; cooperative breeder

Female lions are polyestrous, breeding throughout the year and peaking in the rainy season. Female lions tend to have cubs every 2 years. However, if a female's cubs are killed (usually by an intruding male lion), then the female comes into estrus early and has more cubs. Females are able to breed at 4 years of age and males at 5 years.

One to six cubs are born after a 3.5 month gestation period. There is an interbirth interval of approximately twenty to thirty months. Newborn cubs weigh 1 to 2 kg. Eyes typically open by 11 days, cubs can walk by 15 days and are able to run by 1 month of age. Mother lions keep their cubs in hiding until they reach about 8 weeks of age. The cubs are weaned between 7 and 10 months, however they are dependent upon adults in the pride until they are at least 16 months old.

Breeding interval: Female lions tend to have cubs every 2 years. However, if a female's cubs are killed (usually by an intruding male lion), then she will come into estrus early and have more cubs.

Breeding season: Breeding peaks during the rainy season, but is year-round.

Range number of offspring: 1 to 6.

Average number of offspring: 3.

Average gestation period: 3.5 months.

Range weaning age: 7 to 10 months.

Range time to independence: 16 (low) months.

Average age at sexual or reproductive maturity (female): 4 years.

Average age at sexual or reproductive maturity (male): 5 years.

Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization

Average birth mass: 1300 g.

Average gestation period: 109 days.

Average number of offspring: 3.

Average age at sexual or reproductive maturity (male)
Sex: male:
1095 days.

Average age at sexual or reproductive maturity (female)
Sex: female:
1095 days.

Females are mainly responsible for care of the young. Females nurse their young, but will also nurse the young of their female relatives in the pride if litters are born close together. Cub mortality is lowest when related females in the same pride synchronously reproduce and cross-suckle. Since synchronous reproduction is common in prides, cubs are often raised in crèches where the entire pride helps to raise several litters.

Cubs are often left alone for more than one full day by the time they are 5 to 7 months old. This is a particularly vulnerable time for the cubs to be attacked by predators (often hyenas). Hungry mothers occasionally abandon weak cubs that can not keep up with the pride.

Although males do not directly provide care for the young in a pride, they are important in the protection of the cubs from rival males. So long as a male maintains control over a pride, preventing another male from taking over, the cubs he has sired are at lower risk of infanticide.

Parental Investment: no parental involvement; altricial ; male parental care ; female parental care ; pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Provisioning: Male, Female, Protecting: Male, Female); post-independence association with parents; inherits maternal/paternal territory

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Harrington, E. 2004. "Panthera leo" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html
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Erin Harrington, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Nancy Shefferly, Animal Diversity Web
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Biology

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Lions are the only truly social cats, with related females residing together in prides and related or unrelated males forming coalitions that compete for tenure of prides in fierce and often fatal battles (1). Despite maternal defence, infanticide by the victorious males is common following a pride takeover. This seemingly horrific practice means the lionesses are capable of breeding again sooner, and the reproductive potential of the males is maximised in their often relatively short period of pride tenure. Females are able to breed at four years, males at five, and one to six cubs are born after a 3.5 month gestation period. Females are the predominant care-givers to cubs, which are dependent upon adults until about 16 months old. Related females within a pride are often found to reproduce in synchrony and then cross-suckle their cubs (7). Prides usually consist of four to six adults and their young, which break into smaller groups when hunting (1). Lions are predatory carnivores, with females performing most of the hunting, usually at night to avoid detection (4). They feed upon almost any animal, from rodent to rhino, but medium- to large-sized ungulates, such as antelope, zebra and wildebeest, form the bulk of their prey. They will also scavenge, chasing other predators away from their kills (1).
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Conservation

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In most countries hunting of lions is either prohibited or regulated so that only dangerous animals can be killed, although trophy hunting does remain permissible in a few countries in Africa (6). The Asiatic lion (P. l. persica) is fully protected in India, but another separate population is desperately needed in order to prevent the subspecies being wiped out completely in the wild by an epidemic or other disaster affecting a single population (7) (8). The Palpur-Kuno Wildlife Sanctuary in northern Madhya Pradesh has been identified as a potential reintroduction site in India. In both Africa and Asia, the understanding and cooperation of the community is crucial in ensuring the future of this big cat. Fortunately, the lion is a powerful cultural icon for Africa and one of the highest valued eco-tourism species on the continent, which is now being promoted as an incentive to do everything possible to protect this magnificent 'King of Beasts' (7).
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Description

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It seems that no animal has inspired the imagination of man more than the lion. Characterised as fearsome, courageous and majestic, the lion's strength and ferocity has earned it accolades such as 'King of Beasts' in many cultures (2). As the largest African carnivore and rivalling the tiger (Panthera tigris) as the largest of the 'big cats', the lion is built to prey on animals many times its size, its strong jaws and muscular build emanating an image of sheer power. Males are larger than females and typically posses a mane of hair around their heads, a feature unique amongst the cat family (Felidae) (4). The rest of the coat is short and tawny in colour for both sexes, paler on the underside, without markings. The backs of the ears and the tuft of hair at the tip of the tail are dark brown or black (5). Lion cubs are born with brown rosettes that disappear with maturity, although some lions retain faint spots (6). Asiatic lions (P.l. persica), the only subspecies found outside the African continent, are slightly smaller than their African cousins, and have shorter, thinner manes, clear of the ears, and a fold of skin running the length of their belly that is rare in African lions (7) (8).
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Habitat

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Lions have a broad habitat tolerance, ranging from the savannah woodlands of East Africa to the sands of the Kalahari Desert (2). However, thick brush, scrub, and grass complexes appear to be optimal habitats in providing cover for hunting and denning. Lions have also been recorded to venture high into the mountains of East Africa, up to 4,240 m in Ethiopia's Bale Mountains (4).
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Range

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Formerly ranging throughout Eurasia and Africa (8) (9), lions are now found only in sub-Saharan Africa, and a small isolated population of Asiatic lions (P. l. persica) remains in the Gir Forest in western India (1), where a 2005 census reported just 359 lions (8).
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Status

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Classified as Vulnerable (VU - A2abcd) on the IUCN Red List 2004 (1), and listed on Appendix II of CITES (3). Subspecies: Asiatic lion (Panthera leo persica) classified as Critically Endangered (CR C2a(ii)) on the IUCN Red List 2004 (1), and listed on Appendix I of CITES (3).
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Threats

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Historically lions have been killed for sport (6) and they are generally considered serious problem animals whose existence is in conflict with human settlement and cattle farming (4). The increasing spread of farmlands has reduced the lion's habitat and wild prey base, resulting in increased stock-raiding behaviour (1). This makes lions particularly vulnerable to poisoned carcasses that are put out to eliminate predators. Lions are often seen as vermin and shot on sight, even in protected areas. Publicity of lions as human killers only adds to their unfavourable reputation. Asiatic lions are far more threatened than African lions, and their small gene pool, and therefore genetic instability, puts them at greater risk of epidemics of diseases such as tuberculosis, FIV, and canine distemper virus (6). A population of just 359 individuals is desperately low, but this figure nevertheless represents an increase on a the figures a century ago when rough estimates of the population ranged between 12 and 100 remaining, as a result of intensive hunting (8).
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Description of Panthera leo

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The lion (Panthera leo) is the second-largest living cat after the tiger - with some males exceeding 250 kg (550 lb). Wild lions currently exist in Sub-Saharan Africa and in Asia with an endangered remnant population in Gir Forest National Park in India, having disappeared from North Africa and Southwest Asia in historic times. Until the late Pleistocene, about 10,000 years ago, the lion was the most widespread large land mammal after humans. They were found in most of Africa, across Eurasia from western Europe to India, and in the Americas from the Yukon to Peru. The lion is vulnerable, having seen a possibly irreversible population decline of thirty to fifty percent over the past two decades in its African range. Lion populations are untenable outside designated reserves and national parks. Lions live for ten to fourteen years in the wild, in captivity they may live longer than twenty years. They typically inhabit savanna and grassland, although they may take to bush and forest. Lions are unusually social compared to other cats. A pride of lions consists of related females and offspring and a small number of adult males. Groups of female lions typically hunt together, preying mostly on large ungulates. Lions are apex and keystone predators, although they scavenge as opportunity allows. While lions do not typically hunt humans, some have been known to do so. Highly distinctive, the male lion is easily recognised by its mane, and its face is one of the most widely recognised animal symbols in human culture. Depictions have existed from the Upper Paleolithic period, with carvings and paintings from the Lascaux and Chauvet Caves, through virtually all ancient and medieval cultures where they once occurred. It has been extensively depicted in sculptures, in paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire and have been a key species sought for exhibition in zoos the world over since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.
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BeetleCam Project 2011

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Fact Card

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<Fact Card>


<Physical Description>

<Weight>Males: 330 to 550 pounds; Females: 265 to 395 pounds</Weight>

<Height>3 ½ to 4 feet</Height>

<Length>9-10 feet</Length>

<Lifespan>12-15 years</Lifespan>

<Reproduction>Female lions usually give birth to a litter of 1 to 6 every two years.</Reproduction>

<Geographic Distribution>Southern fringe of Africa’s Sahara desert to northern South Africa but not in areas of moist, tropical forest. A small population survives in NW India as well.</Geographic Distribution>

<Natural Diet>Primarily large animals, such as zebra and wildebeest</Natural Diet>

<Fast Fact>A lion's roar can be heard up to 5.0 miles (8 kilometers) away.</Fast Fact>

</Fact Card>

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Lion Resources from the Field Museum of Natural History

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Explore Lion resources from curator of mammals Bruce Patterson of Chicago’s Field Museum.

Lion Resources

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MammalMAP: the Lion

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Although lions are not the largest, nor the most powerful of all animals, male lions are often referred to as the King of the jungle due to their regal posture. Thery're also one of the 5 big animals, a term coined by hunters of old because of the degree of danger involved in hunting lions. There are many reports, since centuries ago, of hunters injured or killed through the years while trying to hunt lions.

A male lion’s roars can be heard from a distance over 60 km. Lions often roar after a kill or when members of prides are trying to locate each other. Lionesses have a soft short roar that they utter when calling their cubs.

Lions used to occur throughout South Africa, but due to their merciless persecution in South Africa (due to conflicts with humans and their interest), they are now mainly restricted to protected areas. They only occur naturally in the Northern and Eastern parts of South Africa. In the north they mainly occur naturally in the Kruger National Park, and the Private Game reserve farms, bordering Kruger National Park. They also occur naturally in the Kgalagadi Transfrontier Park and in KwaZulu-Natal Parks like Hluhluwe – Imfolozi. Lions have been reintroduced to, amongst others, the Addo National Park, to some provincial parks and reserves, as well as private game reserves throughout South Africa.

They mainly range from woodland and open savanna to desert and arid areas, wherever sufficient prey can be found.

Lions are the largest predators in Africa. They are also the most sociable of the cat family, and they can live in prides of up to 20 and more. Males are territorial and will defend their pride and territory against the intruders, some fights ending in death. There are usually between 1 and 4 adult males (a dominant male and other adults or often brothers of the same litter) per pride, depending on the size of the territory and the size of their pride which consist of females and their cubs, as well as the young males.

Females do most of the hunting, although males do hunt at times. The prides usually hunt together. At any kill of a pride, the dominant will feed first, following other males, and then the females. The cubs will feed last when the others had their fill. Loins hunt by stealth - they stalk their prey or wait in the ambush near the water until the prey animal is close enough to pounce upon. Lions often are opportunistic hunters in the sense that they will kill when the opportunity arise.Lions will also kill any predator if they get a chance and often kill the young of leopards, jackals, hyena, cheetahs, honey badgers, caracal, wild dogs and civets amongst others to naturally eliminate competition for food and territory. They will not always eat these animals but sometimes might.

Lions mate repeatedly (some say every 15 minutes) over a period of 2 to 3 days. They have no specific breeding season. Adult male lions may at times have serious confrontations when battles over a female ensue, at times leading to serious injury or even death for one or both of the lions. Males are much larger and heavier than females, which are of more slender built. Adult males usually have large manes where females do not.


Mass Male: 170 to 235 kg
Female: 130 to 200 kg

Lions are opportunistic hunters. They will eat anything from termites to an elephant if they can. Depending on their territory and species of prey available in the area, they usually hunt medium to large sized animals like wildebeest, impala, zebra, waterbuck, kudu, buffalo and giraffe amongst others and even in some cases hippos, young rhinos and elephants. Giraffe have been known to inflict serious injuries to lions and often kill lions, while defending themselves by kicking at the lions especially with hind legs.

Lions are often found following large or smaller herds of buffalo while waiting for chances to catch and kill one, although buffaloes are not that easy to catch and kill due to the fact that they will not always run away – they will group together (often forming a circle around the young) and try to stand their ground in an effort to defend themselves and their young ones against the lions. Lions are also often injured or killed by buffaloes, especially by the bulls, which can cause a lot of damage with their horns as well as by stomping on the lions with their hard hooves.

For more information on MammalMAP, visit the MammalMAPvirtual museumorblog.

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One Species at a Time Podcast: Lions

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Does the mane really make the lion? Certainly, luxurious locks are the feature that sets Panthera leo apart from the other large cats. But surprisingly, not all male lions have manes. And manes used to cover more of the lion than just the head. Ari Daniel Shapiro speaks with archivist Connie Rinaldo of Harvard University and curator of mammals Bruce Patterson of Chicago’s Field Museum to learn about the diversity of lions in the distant past and the challenges they face in the present.

Listen to this podcast

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Pleistocene Re-wilding

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This species is one of a number which have been included in various “Pleistocene rewilding” plans. Pleistocene rewilding is the proposed practice of restoring ecosystems to their state in the Pleistocene, roughly 10,000 years ago. This contrasts the standard conservation benchmark, particularly in North America, of restoring ecosystems to their pre-Columbian or pre-industrial state. In both Eurasia and North America, the Pleistocene was characterized by much greater diversity and numbers of large herbivores and predators, including proboscidians, equids, camelids, and felidae (Donlan et al 2006; Zimov 2005). The process of restoration would involve the reintroduction of extant species in their historic range, as well as the introduction of ‘proxy organisms’ to replace the ecological functionality of extinct organisms (Donlan et al 2006).

There are three central theoretical goals to Pleistocene rewilding. In Siberia, a team led by Sergey Zimov is investigating the role of large herbivores as ecosystem engineers. It is thought that herbivory pressure could play a central role in maintaining a grass-dominated plant community, as opposed to either tree- or moss-dominated. Grasslands are known to be more stable carbon sinks than either mossy or forested tundra, due to the rapidity of their biogeochemical cycling (Zimov 2005). In principle, then, reintroducing Pleistocene fauna could have positive climate change mitigation effects. Proposals in North America have focused instead on the preservation of ecological dynamics. Proponents of Pleistocene rewilding argue that due to the strong ecological interactions of megafauna, it is likely that their extinction at the end of the Pleistocene would have caused cascading ecological disruptions lasting until the present time (Donlan et al 2006). Additionally, introduction programs could provide a new lease on life for extant, endangered megafauna species, such as cheetahs and Asian elephants (Rubenstein 2006).

Pleistocene rewilding, while headline-grabbing, is by no means the standard of modern conservation biology. There are a number of objections to the proposals of Pleistocene rewilders, summarized by Rubenstein et al (2006). The introduction of species which have been locally extinct for thousands of years, and more particularly the introduction of modern relatives of extinct species, carries many risks: the potential for invasive species, catastrophic disruption of existing ecosystems, inadvertent introduction of disease organisms, and unpredictable behavior of introduced species. Additionally, while paleoecology is a growing field, there is still a fair amount of uncertainty about the actual ecosystem functions of the Pleistocene.

Species which Zimov and his colleagues in Siberia are experimenting with bison, musk oxen, Przewalski’s horse, and Siberian tigers (Zimov 2005). Small-scale introductions have already begun in Yakutia. Donlan et al propose introducing Przewalski’s horse, Bolson tortoises, Bactrian camels, cheetahs, lions, and elephants into the Western United States (Donlan et al 2005). While some individuals of these species are present on privately owned land, there are no free-living populations in North America at this time.

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lion - Bing Images

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Some images of lions. I hope people like them!

Lion

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The lion (Panthera leo) is a large cat of the genus Panthera native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adult male lions are larger than females and have a prominent mane. It is a social species, forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator; although some lions scavenge when opportunities occur and have been known to hunt humans, lions typically do not actively seek out and prey on humans.

The lion inhabits grasslands, savannahs and shrublands. It is usually more diurnal than other wild cats, but when persecuted, it adapts to being active at night and at twilight. During the Neolithic period, the lion ranged throughout Africa and Eurasia from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in western India. It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have declined by about 43% since the early 1990s. Lion populations are untenable outside designated protected areas. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes for concern.

One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire and have been a key species sought for exhibition in zoological gardens across the world since the late 18th century. Cultural depictions of lions were prominent in Ancient Egypt, and depictions have occurred in virtually all ancient and medieval cultures in the lion's historic and current range.

Etymology

The English word lion is derived via Anglo-Norman liun from Latin leōnem (nominative: leō), which in turn was a borrowing from Ancient Greek λέων léōn. The Hebrew word לָבִיא lavi may also be related.[4] The generic name Panthera is traceable to the classical Latin word 'panthēra' and the ancient Greek word πάνθηρ 'panther'.[5]

Taxonomy

The upper cladogram is based on the 2006 study,[6][7] the lower one on the 2010[8] and 2011[9] studies.

Felis leo was the scientific name used by Carl Linnaeus in 1758, who described the lion in his work Systema Naturae.[3] The genus name Panthera was coined by Lorenz Oken in 1816.[10] Between the mid-18th and mid-20th centuries, 26 lion specimens were described and proposed as subspecies, of which 11 were recognised as valid in 2005.[1] They were distinguished mostly by the size and colour of their manes and skins.[11]

Subspecies

Range map showing distribution of subspecies and clades

In the 19th and 20th centuries, several lion type specimens were described and proposed as subspecies, with about a dozen recognised as valid taxa until 2017.[1] Between 2008 and 2016, IUCN Red List assessors used only two subspecific names: P. l. leo for African lion populations, and P. l. persica for the Asiatic lion population.[2][12][13] In 2017, the Cat Classification Task Force of the Cat Specialist Group revised lion taxonomy, and recognises two subspecies based on results of several phylogeographic studies on lion evolution, namely:[14]

  • P. l. leo (Linnaeus, 1758) − the nominate lion subspecies includes the Asiatic lion, the regionally extinct Barbary lion, and lion populations in West and northern parts of Central Africa.[14] Synonyms include P. l. persica (Meyer, 1826), P. l. senegalensis (Meyer, 1826), P. l. kamptzi (Matschie, 1900), and P. l. azandica (Allen, 1924).[1] Multiple authors referred to it as 'northern lion' and 'northern subspecies'.[15][16]
  • P. l. melanochaita (Smith, 1842) − includes the extinct Cape lion and lion populations in East and Southern African regions.[14] Synonyms include P. l. somaliensis (Noack 1891), P. l. massaica (Neumann, 1900), P. l. sabakiensis (Lönnberg, 1910), P. l. bleyenberghi (Lönnberg, 1914), P. l. roosevelti (Heller, 1914), P. l. nyanzae (Heller, 1914), P. l. hollisteri (Allen, 1924), P. l. krugeri (Roberts, 1929), P. l. vernayi (Roberts, 1948), and P. l. webbiensis (Zukowsky, 1964).[1][11] It has been referred to as 'southern subspecies' and 'southern lion'.[16]

However, there seems to be some degree of overlap between both groups in northern Central Africa. DNA analysis from a more recent study indicates, that Central African lions are derived from both northern and southern lions, as they cluster with P. leo leo in mtDNA-based phylogenies whereas their genomic DNA indicates a closer relationship with P. leo melanochaita.[17]

Lion samples from some parts of the Ethiopian Highlands cluster genetically with those from Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East Africa. Researchers therefore assume Ethiopia is a contact zone between the two subspecies.[18] Genome-wide data of a wild-born historical lion sample from Sudan showed that it clustered with P. l. leo in mtDNA-based phylogenies, but with a high affinity to P. l. melanochaita. This result suggested that the taxonomic position of lions in Central Africa may require revision.[19]

Fossil records

Skull of an American lion on display at the National Museum of Natural History

Other lion subspecies or sister species to the modern lion existed in prehistoric times:[20]

Evolution

red Panthera spelaea
blue Panthera atrox
green Panthera leo

Maximal range of the modern lion
and its prehistoric relatives
in the late Pleistocene

The Panthera lineage is estimated to have genetically diverged from the common ancestor of the Felidae around 9.32 to 4.47 million years ago to 11.75 to 0.97 million years ago,[6][33][34] and the geographic origin of the genus is most likely northern Central Asia.[35] Results of analyses differ in the phylogenetic relationship of the lion; it was thought to form a sister group with the jaguar (P. onca) that diverged 3.46 to 1.22 million years ago,[6] but also with the leopard (P. pardus) that diverged 3.1 to 1.95 million years ago[8][9] to 4.32 to 0.02 million years ago. Hybridisation between lion and snow leopard (P. uncia) ancestors possibly continued until about 2.1 million years ago.[34] The lion-leopard clade was distributed in the Asian and African Palearctic since at least the early Pliocene.[35] The earliest fossils recognisable as lions were found at Olduvai Gorge in Tanzania and are estimated to be up to 2 million years old.[33]

Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on mutation rate per generation time of the modern lion. There is no evidence for gene flow between the two lineages, indicating that they did not share the same geographic area.[19] The Eurasian and American cave lions became extinct at the end of the last glacial period without mitochondrial descendants on other continents.[27][36][37] The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago.[38] They shared a common ancestor probably between 98,000 and 52,000 years ago.[19] Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As the rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia between 38,800 and 8,300 years ago.[38]

Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa. Genetic evidence revealed numerous mutations in lion samples from East and Southern Africa, which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa.[39] A whole genome-wide sequence of lion samples showed that samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated, possibly migrating through corridors in the Nile Basin during the early Holocene.[19]

Hybrids

In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose.[40][41] The liger is bigger than a lion and a tiger, whereas most tigons are relatively small compared to their parents because of reciprocal gene effects.[42][43] The leopon is a hybrid between a lion and leopard.[44]

Description

A tuft at the end of the tail is a distinct characteristic of the lion.
Skeleton

The lion is a muscular, broad-chested cat with a short, rounded head, a reduced neck and round ears; males have broader heads. The fur varies in colour from light buff to silvery grey, yellowish red and dark brown. The colours of the underparts are generally lighter. A new-born lion has dark spots, which fade as the cub reaches adulthood, although faint spots often may still be seen on the legs and underparts.[45][46] The tail of all lions ends in a dark, hairy tuft that in some lions conceals an approximately 5 mm (0.20 in)-long, hard "spine" or "spur" that is formed from the final, fused sections of tail bone. The functions of the spur are unknown. The tuft is absent at birth and develops at around 5+12 months of age. It is readily identifiable by the age of seven months.[47]

Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened, and has a slightly shorter postorbital region and broader nasal openings than those of the tiger. Due to the amount of skull variation in the two species, usually only the structure of the lower jaw can be used as a reliable indicator of species.[48][49]

Skeletal muscles of the lion make up 58.8% of its body weight and represents the highest percentage of muscles among mammals.[50][51]

Size

Among felids, the lion is second only to the tiger in size.[46] The size and weight of adult lions varies across its range and habitats.[52][53][54][55] Accounts of a few individuals that were larger than average exist from Africa and India.[45][56][57][58]

Mane

A six-year-old male in Phinda Private Game Reserve
Male in Pendjari National Park

The male lion's mane is the most recognisable feature of the species.[11] It may have evolved around 320,000–190,000 years ago.[60] It grows downwards and backwards covering most of the head, neck, shoulders, and chest. The mane is typically brownish and tinged with yellow, rust and black hairs.[46] It starts growing when lions enter adolescence, when testosterone levels increase, and reach their full size at around four years old.[61] Cool ambient temperatures in European and North American zoos may result in a heavier mane.[62] Asiatic lions usually have sparser manes than average African lions.[63]

This feature likely evolved to signal the fitness of males to females and not to protect the neck. During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back and hindquarters. Injured lions begin to lose their manes, further evidence against it being protective. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hairs with higher testosterone but are also more vulnerable to heat stress. Unlike in other felid species, female lions consistently interact with multiple males at once, hence why the mane did not evolve in other male cats.[64]

Almost all male lions in Pendjari National Park are either maneless or have very short manes.[65] Maneless lions have also been reported in Senegal, in Sudan's Dinder National Park and in Tsavo East National Park, Kenya.[66] Castrated lions often have little to no mane because the removal of the gonads inhibits testosterone production.[67] Increased testosterone may be the cause of maned lionesses reported in northern Botswana.[68]

Colour variation

The white lion is a rare morph with a genetic condition called leucism which is caused by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool. Nevertheless, 17 births have been recorded in five prides between 2007 and 2015.[69] White lions are selected for breeding in captivity.[70] They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts.[71]

Distribution and habitat

African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and savannahs, scrub bordering rivers and open woodlands with bushes. It rarely enters closed forests. On Mount Elgon, the lion has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.[45] Savannahs with an annual rainfall of 300 to 1,500 mm (12 to 59 in) make up the majority of lion habitat in Africa, estimated at 3,390,821 km2 (1,309,203 sq mi) at most, but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa.[72] The Asiatic lion now survives only in and around Gir National Park in Gujarat, western India. Its habitat is a mixture of dry savannah forest and very dry, deciduous scrub forest.[12]

Historical range

In Africa, the range of the lion originally spanned most of the central African rainforest zone and the Sahara desert.[73] In the 1960s, it became extinct in North Africa, except in the southern part of Sudan.[74][72][75]

In southern Europe and Asia, the lion once ranged in regions where climatic conditions supported an abundance of prey.[76] In Greece, it was common as reported by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100.[45] It was present in the Caucasus until the 10th century.[49] It lived in Palestine until the Middle Ages, and in Southwest Asia until the late 19th century. By the late 19th century, it had been extirpated in most of Turkey.[77] The last live lion in Iran was sighted in 1942 about 65 km (40 mi) northwest of Dezful,[78] although the corpse of a lioness was found on the banks of the Karun river in Khūzestān Province in 1944.[79] It once ranged from Sind and Punjab in Pakistan to Bengal and the Narmada River in central India.[80]

Behaviour and ecology

Lions spend much of their time resting; they are inactive for about twenty hours per day.[81] Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, grooming and defecating. Intermittent bursts of activity continue until dawn, when hunting most often takes place. They spend an average of two hours a day walking and fifty minutes eating.[82]

Group organisation

Lion pride in Etosha National Park
A lioness (left) and two males in Masai Mara

The lion is the most social of all wild felid species, living in groups of related individuals with their offspring. Such a group is called a "pride". Groups of male lions are called "coalitions".[83] Females form the stable social unit in a pride and do not tolerate outside females.[84] The majority of females remain in their birth prides while all males and some females will disperse.[85] The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have been observed.[86] The sole exception to this pattern is the Tsavo lion pride that always has just one adult male.[87] Prides act as fission–fusion societies, and members will split into subgroups which keep in contact with roars.[88]

Nomadic lions range widely and move around sporadically, either in pairs or alone.[83] Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.[89] Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them.[90] Males spend years in a nomadic phase before gaining residence in a pride.[91] A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.[92] In Kruger National Park, dispersing male lions move more than 25 km (16 mi) away from their natal pride in search of their own territory. Female lions stay closer to their natal pride. Therefore, female lions in an area are more closely related to each other than male lions in the same area.[93]

The evolution of sociability in lions was likely driven both by high population density and the clumped resources of savannah habitats. The larger the pride, the more high-quality territory they can defend; "hotspots" being near river confluences, where the cats have better access to water, prey and shelter (via vegetation).[94][95] The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".[83] Males associated with a pride patrol the fringes.[46] Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defense against intruders, while others lag behind.[96] Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.[97] Alternatively, there may be rewards associated with being a leader that fends off intruders; the rank of lionesses in the pride is reflected in these responses.[98] The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.[89] Dominance hierarchies do not appear to exist among individuals of either sex in a pride.[99]

Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up to three males, forming a loose pride while females associate with up to 12 other females, forming a stronger pride together with their cubs. Female and male lions associate only when mating.[100] Coalitions of males hold territory for a longer time than single lions. Males in coalitions of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the others and mates more frequently.[101]

Hunting and diet

Male lion and cub feeding on a Cape buffalo in Sabi Sand Game Reserve
Four lionesses catching a buffalo in the Serengeti
A skeletal mount of a lion attacking a common eland, on display at The Museum of Osteology

The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum.[102][103] Its prey consists mainly of ungulates weighing 190–550 kg (420–1,210 lb), particularly blue wildebeest, plains zebra, African buffalo, gemsbok and giraffe. They also hunt common warthog depending on availability, despite weighing less than the preferred weight range.[104] In India, chital and sambar deer are the most common wild prey,[46][104][105] while livestock contributes significantly to lion kills outside protected areas.[106] They usually avoid fully grown adult elephants, rhinoceroses and hippopotamus and small prey like dik-dik, hyraxes, hares and monkeys.[104][107] Unusual prey include porcupines and small reptiles. Lions kill other predators but seldom consume them.[108]

Young lions first display stalking behaviour at around three months of age, although they do not participate in hunting until they are almost a year old and begin to hunt effectively when nearing the age of two.[109] Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.[89] In Chobe National Park, large prides have been observed hunting African bush elephants up to around 15 years old in exceptional cases, with the victims being calves, juveniles, and even subadults.[110][111] In typical hunts, each lioness has a favoured position in the group, either stalking prey on the "wing", then attacking, or moving a smaller distance in the centre of the group and capturing prey fleeing from other lionesses. Males attached to prides do not usually participate in group hunting.[112] Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.[113] They may join in the hunting of large, slower-moving prey like buffalo; and even hunt them on their own. Moderately-sized hunting groups generally have higher success rates than lone females and larger groups.[114]

Lions are not particularly known for their stamina. For instance, a lioness's heart comprises only 0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's heart comprises almost 1% of its body weight.[115] Thus, lions run quickly only in short bursts at about 48–59 km/h (30–37 mph) and need to be close to their prey before starting the attack.[116] One study in 2018 recorded a lion running at a top speed of 74.1 km/h (46.0 mph).[117] They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[118] The lion's attack is short and powerful; they attempt to catch prey with a fast rush and final leap. They usually pull it down by the rump and kill by a clamping bite to the throat or muzzle.[119]

Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover.[120] They tend to squabble over kills, particularly the males. Cubs suffer most when food is scarce but otherwise all pride members eat their fill, including old and crippled lions, which can live on leftovers.[89] Large kills are shared more widely among pride members.[121] An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb).[122] Lions gorge themselves and eat up to 30 kg (66 lb) in one session.[79] If it is unable to consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride retreats to shade with one or two males standing guard.[120] Lions defend their kills from scavengers such as vultures and hyenas.[89]

Lions scavenge on carrion when the opportunity arises, scavenging animals dead from natural causes such as disease or those that were killed by other predators. Scavenging lions keep a constant lookout for circling vultures, which indicate the death or distress of an animal.[123] Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions.[55] Carrion is thought to provide a large part of lion diet.[124]

Predatory competition

Lion attacked by spotted hyenas in Sabi Sand Game Reserve
Lioness stealing a kill from a leopard in Kruger National Park

Lions and spotted hyenas occupy a similar ecological niche and compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%.[125] Lions typically ignore hyenas unless they are on a kill or are being harassed, while the latter tend to visibly react to the presence of lions with or without the presence of food. In the Ngorongoro crater, lions subsist largely on kills stolen from hyenas, causing them to increase their kill rate.[126] In Botswana's Chobe National Park, the situation is reversed as hyenas there frequently challenge lions and steal their kills, obtaining food from 63% of all lion kills.[127] When confronted on a kill, hyenas may either leave or wait patiently at a distance of 30–100 m (98–328 ft) until the lions have finished.[128] Hyenas feed alongside lions and force them off a kill. The two species attack one another even when there is no food involved for no apparent reason.[129] Lions can account for up to 71% of hyena deaths in Etosha National Park. Hyenas have adapted by frequently mobbing lions that enter their home ranges.[130] When the lion population in Kenya's Masai Mara National Reserve declined, the spotted hyena population increased rapidly.[131]

Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults when given the chance.[132] Cheetahs often lose their kills to lions or other predators.[133] A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.[134] Cheetahs avoid their competitors by hunting at different times and habitats.[135] Leopards take refuge in trees, but lionesses occasionally attempt to climb up and retrieve their kills.[136]

Lions similarly dominate African wild dogs, taking their kills and slaying pups or adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.[137] However, there are a few reported cases of old and wounded lion falling prey to wild dogs.[138][139] Nile crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.[140]

Reproduction and life cycle

Lions mating at Masai Mara
A lion cub in Masai Mara

Most lionesses reproduce by the time they are four years of age.[141] Lions do not mate at a specific time of year and the females are polyestrous.[142] Like those of other cats, the male lion's penis has spines that point backward. During withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation.[143][144] A lioness may mate with more than one male when she is in heat.[145] Lions of both sexes may be involved in group homosexual and courtship activities. Males will also head-rub and roll around with each other before simulating sex together.[146][147] Generation length of the lion is about seven years.[148] The average gestation period is around 110 days;[142] the female gives birth to a litter of between one and four cubs in a secluded den, which may be a thicket, a reed-bed, a cave, or some other sheltered area, usually away from the pride. She will often hunt alone while the cubs are still helpless, staying relatively close to the den.[149] Lion cubs are born blind, their eyes opening around seven days after birth. They weigh 1.2–2.1 kg (2.6–4.6 lb) at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.[150] To avoid a buildup of scent attracting the attention of predators, the lioness moves her cubs to a new den site several times a month, carrying them one-by-one by the nape of the neck.[149]

Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.[149] Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time.[89][151] When first introduced to the rest of the pride, lion cubs lack confidence when confronted with adults other than their mother. They soon begin to immerse themselves in the pride life, however, playing among themselves or attempting to initiate play with the adults.[151] Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. Male tolerance of the cubs varies—one male could patiently let the cubs play with his tail or his mane, while another may snarl and bat the cubs away.[152]

Video of a lioness and her cubs in Phinda Reserve

Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young, which suckle indiscriminately from any or all of the nursing females in the pride. The synchronisation of births is advantageous because the cubs grow to being roughly the same size and have an equal chance of survival, and sucklings are not dominated by older cubs.[89][151] Weaning occurs after six or seven months. Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride. They begin to age and weaken at between 10 and 15 years of age at the latest.[153]

When one or more new males oust the previous males associated with a pride, the victors often kill any existing young cubs, perhaps because females do not become fertile and receptive until their cubs mature or die. Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male.[154] Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age,[155] while some females may leave when they reach the age of two.[85] When a new male lion takes over a pride, adolescents both male and female may be evicted.[156]

Health and mortality

Lions in a tree near Lake Nakuru

Lions may live 12–17 years.[46] Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.[157] Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at a kill.[158] Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants. Careless lions may be maimed when hunting prey.[159]

Ticks commonly infest the ears, neck and groin regions of the lions.[160][161] Adult forms of several tapeworm species of the genus Taenia have been isolated from lion intestines, having been ingested as larvae in antelope meat.[162] Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (Stomoxys calcitrans) in 1962, resulting in lions becoming emaciated and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals.[163] A more recent outbreak in 2001 killed six lions.[164]

Captive lions have been infected with canine distemper virus (CDV) since at least the mid-1970s.[165] CDV is spread by domestic dogs and other carnivores; a 1994 outbreak in Serengeti National Park resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from pneumonia and encephalitis.[166] Feline immunodeficiency virus and lentivirus also affect captive lions.[167][168]

Communication

Head rubbing among pride members is a common social behaviour.

When resting, lion socialisation occurs through a number of behaviours; the animal's expressive movements are highly developed. The most common peaceful, tactile gestures are head rubbing and social licking,[169] which have been compared with the role of allogrooming among primates.[170] Head rubbing—nuzzling the forehead, face and neck against another lion—appears to be a form of greeting[171] and is seen often after an animal has been apart from others or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.[172] Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked; this behaviour may have arisen out of utility because lions cannot lick these areas themselves.[173]

A captive lion roaring

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Lions have an array of facial expressions and body postures that serve as visual gestures.[174] A common facial expression is the "grimace face" or flehmen response, which a lion makes when sniffing chemical signals and involves an open mouth with bared teeth, raised muzzle, wrinkled nose closed eyes and relaxed ears.[175] Lions also use chemical and visual marking; males will spray and scrape plots of ground and objects within the territory.[174]

The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of growling, snarling, meowing and roaring. Other sounds produced include purring, puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of 8 kilometres (5 mi).[176] They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into a series of shorter ones.[177][178]

Conservation

The lion is listed as Vulnerable on the IUCN Red List. The Indian population is listed on CITES Appendix I and the African population on CITES Appendix II.[2]

In Africa

Video of a wild lioness

Several large and well-managed protected areas in Africa host large lion populations. Where an infrastructure for wildlife tourism has been developed, cash revenue for park management and local communities is a strong incentive for lion conservation.[2] Most lions now live in East and Southern Africa; their numbers are rapidly decreasing, and fell by an estimated 30–50% in the late half of the 20th century. Primary causes of the decline include disease and human interference.[2] In 1975, it was estimated that since the 1950s, lion numbers had decreased by half to 200,000 or fewer.[179] Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.[180][74]

In the Republic of the Congo, Odzala-Kokoua National Park was considered a lion stronghold in the 1990s. By 2014, no lions were recorded in the protected area so the population is considered locally extinct.[181] The West African lion population is isolated from the one in Central Africa, with little or no exchange of breeding individuals. In 2015, it was estimated that this population consists of about 400 animals, including fewer than 250 mature individuals. They persist in three protected areas in the region, mostly in one population in the W A P protected area complex, shared by Benin, Burkina Faso and Niger. This population is listed as Critically Endangered.[13] Field surveys in the WAP ecosystem revealed that lion occupancy is lowest in the W National Park, and higher in areas with permanent staff and thus better protection.[182]

A population occurs in Cameroon's Waza National Park, where between approximately 14 and 21 animals persisted as of 2009.[183] In addition, 50 to 150 lions are estimated to be present in Burkina Faso's Arly-Singou ecosystem.[184] In 2015, an adult male lion and a female lion were sighted in Ghana's Mole National Park. These were the first sightings of lions in the country in 39 years.[185] In the same year, a population of up to 200 lions that was previously thought to have been extirpated was filmed in the Alatash National Park, Ethiopia, close to the Sudanese border.[186][187]

In 2005, Lion Conservation Strategies were developed for West and Central Africa, and or East and Southern Africa. The strategies seek to maintain suitable habitat, ensure a sufficient wild prey base for lions, reduce factors that lead to further fragmentation of populations, and make lion–human coexistence sustainable.[188][189] Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating human–lion conflict.[190]

In Asia

A lioness in Gir National Park

The last refuge of the Asiatic lion population is the 1,412 km2 (545 sq mi) Gir National Park and surrounding areas in the region of Saurashtra or Kathiawar Peninsula in Gujarat State, India. The population has risen from approximately 180 lions in 1974 to about 400 in 2010.[191] It is geographically isolated, which can lead to inbreeding and reduced genetic diversity. Since 2008, the Asiatic lion has been listed as Endangered on the IUCN Red List.[12] By 2015, the population had grown to 523 individuals inhabiting an area of 7,000 km2 (2,700 sq mi) in Saurashtra.[192][193][194] The Asiatic Lion Census conducted in 2017 recorded about 650 individuals.[195]

The presence of numerous human habitations close to the National Park results in conflict between lions, local people and their livestock.[196][192] Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check.[197] The establishment of a second, independent Asiatic lion population in Kuno Wildlife Sanctuary, located in Madhya Pradesh was planned but in 2017, the Asiatic Lion Reintroduction Project seemed unlikely to be implemented.[198][199]

Captive breeding

Two captive male Asiatic lions in Sanjay Gandhi National Park, India

Lions imported to Europe before the middle of the 19th century were possibly foremost Barbary lions from North Africa, or Cape lions from Southern Africa.[200] Another 11 animals thought to be Barbary lions kept in Addis Ababa Zoo are descendants of animals owned by Emperor Haile Selassie. WildLink International in collaboration with Oxford University launched an ambitious International Barbary Lion Project with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the Atlas Mountains of Morocco.[201] However, a genetic analysis showed that the captive lions at Addis Ababa Zoo were not Barbary lions, but rather closely related to wild lions in Chad and Cameroon.[202]

In 1982, the Association of Zoos and Aquariums started a Species Survival Plan for the Asiatic lion to increase its chances of survival. In 1987, it was found that most lions in North American zoos were hybrids between African and Asiatic lions.[203] Breeding programs need to note origins of the participating animals to avoid cross-breeding different subspecies and thus reducing their conservation value.[204] Captive breeding of lions was halted to eliminate individuals of unknown origin and pedigree. Wild-born lions were imported to American zoos from Africa between 1989 and 1995. Breeding was continued in 1998 in the frame of an African lion Species Survival Plan.[205]

About 77% of the captive lions registered in the International Species Information System in 2006 were of unknown origin; these animals might have carried genes that are extinct in the wild and may therefore be important to the maintenance of the overall genetic variability of the lion.[62]

Interactions with humans

In zoos and circuses

Lion at Melbourne Zoo
19th-century etching of a lion tamer in a cage with lions and tigers

Lions are part of a group of exotic animals that have been central to zoo exhibits since the late 18th century. Although many modern zoos are more selective about their exhibits,[206] there are more than 1,000 African and 100 Asiatic lions in zoos and wildlife parks around the world. They are considered an ambassador species and are kept for tourism, education and conservation purposes.[207] Lions can live over twenty years in captivity; for example, three sibling lions at the Honolulu Zoo lived to the age of 22 in 2007.[208][209]

The first European "zoos" spread among noble and royal families in the 13th century, and until the 17th century were called seraglios. At that time, they came to be called menageries, an extension of the cabinet of curiosities. They spread from France and Italy during the Renaissance to the rest of Europe.[210] In England, although the seraglio tradition was less developed, lions were kept at the Tower of London in a seraglio established by King John in the 13th century;[211][212] this was probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in Woodstock, Oxfordshire, where according to William of Malmesbury lions had been stocked.[213]

Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.[214] Further changes took place in the early 20th century when Carl Hagenbeck designed enclosures with concrete "rocks", more open space and a moat instead of bars, more closely resembling a natural habitat. Hagenbeck designed lion enclosures for both Melbourne Zoo and Sydney's Taronga Zoo; although his designs were popular, the use of bars and caged enclosures prevailed in many zoos until the 1960s.[215] In the late 20th century, larger, more natural enclosures and the use of wire mesh or laminated glass instead of lowered dens allowed visitors to come closer than ever to the animals; some attractions such as the Cat Forest/Lion Overlook of Oklahoma City Zoological Park placed the den on ground level, higher than visitors.[216]

Lion taming has been part of both established circuses and individual acts such as Siegfried & Roy. The practice began in the early 19th century by Frenchman Henri Martin and American Isaac Van Amburgh, who both toured widely and whose techniques were copied by a number of followers.[217] Van Amburgh performed before Queen Victoria in 1838 when he toured Great Britain. Martin composed a pantomime titled Les Lions de Mysore ("the lions of Mysore"), an idea Amburgh quickly borrowed. These acts eclipsed equestrianism acts as the central display of circus shows and entered public consciousness in the early 20th century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.[217] The ultimate proof of a tamer's dominance and control over a lion is demonstrated by the placing of the tamer's head in the lion's mouth. The now-iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[218]

Hunting and games

Bas-relief of a wounded lioness from Nineveh, c. 645–635 BC

Lion hunting has occurred since ancient times and was often a royal tradition, intended to demonstrate the power of the king over nature. Such hunts took place in a reserved area in front of an audience. The monarch was accompanied by his men and controls were put in place to increase their safety and ease of killing. The earliest surviving record of lion hunting is an ancient Egyptian inscription dated circa 1380 BC that mentions Pharaoh Amenhotep III killing 102 lions in ten years "with his own arrows". The Assyrian emperor Ashurbanipal had one of his lion hunts depicted on a sequence of Assyrian palace reliefs c. 640 BC, known as the Lion Hunt of Ashurbanipal. Lions were also hunted during the Mughal Empire, where Emperor Jahangir is said to have excelled at it.[219] In Ancient Rome, lions were kept by emperors for hunts, gladiator fights and executions.[220]

The Maasai people have traditionally viewed the killing of lions as a rite of passage. Historically, lions were hunted by individuals, however, due to reduced lion populations, elders discourage solo lion hunts.[221] During the European colonisation of Africa in the 19th century, the hunting of lions was encouraged because they were considered pests and lion skins were sold for £1 each.[222] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[223] Trophy hunting of lions in recent years has been met with controversy, notably with the killing of Cecil the lion in mid-2015.[224]

Man-eating

The Tsavo maneaters of East Africa on display in the Field Museum of Natural History in Chicago

Lions do not usually hunt humans but some (usually males) seem to seek them out. One well-publicised case is the Tsavo maneaters; in 1898, 28 officially recorded railway workers building the Uganda Railway were taken by lions over nine months during the construction of a bridge in Kenya.[225] The hunter who killed the lions wrote a book detailing the animals' predatory behaviour; they were larger than normal and lacked manes, and one seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favoured by all researchers; an analysis of teeth and jaws of man-eating lions in museum collections suggests that while tooth decay may explain some incidents, prey depletion in human-dominated areas is a more likely cause of lion predation on humans.[226] Sick or injured animals may be more prone to man-eating but the behaviour is not unusual, nor necessarily aberrant.[227]

Lions' proclivity for man-eating has been systematically examined. American and Tanzanian scientists report that man-eating behaviour in rural areas of Tanzania increased greatly from 1990 to 2005. At least 563 villagers were attacked and many eaten over this period. The incidents occurred near Selous Game Reserve in Rufiji River and in Lindi Region near the Mozambican border. While the expansion of villages into bush country is one concern, the authors argue conservation policy must mitigate the danger because in this case, conservation contributes directly to human deaths. Cases in Lindi in which lions seize humans from the centres of substantial villages have been documented.[228] Another study of 1,000 people attacked by lions in southern Tanzania between 1988 and 2009 found that the weeks following the full moon, when there was less moonlight, were a strong indicator of increased night-time attacks on people.[229]

According to Robert R. Frump, Mozambican refugees regularly crossing Kruger National Park, South Africa, at night are attacked and eaten by lions; park officials have said man-eating is a problem there. Frump said thousands may have been killed in the decades after apartheid sealed the park and forced refugees to cross the park at night. For nearly a century before the border was sealed, Mozambicans had regularly crossed the park in daytime with little harm.[230]

Cultural significance

Lions carved on a rock weight, Jiroft culture, Iran, 3rd mil. BC

The lion is one of the most widely recognised animal symbols in human culture. It has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature.[45] It appeared as a symbol for strength and nobility in cultures across Europe, Asia and Africa, despite incidents of attacks on people. The lion has been depicted as "king of the jungle" and "king of beasts", and thus became a popular symbol for royalty and stateliness.[231] The lion is also used as a symbol of sporting teams.[232]

Africa

Granite statue of the Egyptian goddess Sekhmet from the Luxor Temple, dated 1403–1365 BC, exhibited in the National Museum of Denmark

In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts.[233] In some cultures, the lion symbolises power and royalty.[234] In the Swahili language, the lion is known as simba which also means "aggressive", "king" and "strong".[54] Some rulers had the word "lion" in their nickname. Sundiata Keita of the Mali Empire was called "Lion of Mali".[235] The founder of the Waalo kingdom is said to have been raised by lions and returned to his people part-lion to unite them using the knowledge he learned from the lions.[234]

In parts of West Africa, lions symbolised the top class of their social hierarchies.[234] In more heavily forested areas where lions were rare, the leopard represented the top of the hierarchy.[233] In parts of West and East Africa, the lion is associated with healing and provides the connection between seers and the supernatural. In other East African traditions, the lion represents laziness.[234] In much of African folklore, the lion is portrayed as having low intelligence and is easily tricked by other animals.[235]

The ancient Egyptians portrayed several of their war deities as lionesses, which they revered as fierce hunters. Egyptian deities associated with lions include Sekhmet, Bast, Mafdet, Menhit, Pakhet and Tefnut.[231] These deities were often connected with the sun god Ra and his fierce heat, and their dangerous power was invoked to guard people or sacred places. The sphinx, a figure with a lion's body and the head of a human or other creature, represented a pharaoh or deity who had taken on this protective role.[236]

Asia

Roaring and striding lion from the Throne Room of Nebuchadnezzar II, 6th century BC, from Babylon, Iraq

The lion was a prominent symbol in ancient Mesopotamia from Sumer up to Assyrian and Babylonian times, where it was strongly associated with kingship.[237] Lions were among the major symbols of the goddess Inanna/Ishtar.[238][239] The Lion of Babylon was the foremost symbol of the Babylonian Empire.[240] The Lion of Judah is the biblical emblem of the tribe of Judah and the later Kingdom of Judah.[241] Lions are frequently mentioned in the Bible, notably in the Book of Daniel, in which the eponymous hero refuses to worship King Darius and is forced to sleep in the lions' den where he is miraculously unharmed (Dan 6). In the Book of Judges, Samson kills a lion as he travels to visit a Philistine woman.(Judg 14).[242]

Indo-Persian chroniclers regarded the lion as keeper of order in the realm of animals. The Sanskrit word mrigendra signifies a lion as king of animals in general or deer in particular.[243] Narasimha, the man-lion, is one of ten avatars of the Hindu god Vishnu.[244] Singh is an ancient Indian vedic name meaning "lion", dating back over 2,000 years. It was originally used only by Rajputs, a Hindu Kshatriya or military caste but is used by millions of Hindu Rajputs and more than twenty million Sikhs today.[245] The Lion Capital of Ashoka, erected by Emperor Ashoka in the 3rd century CE, depicts four lions standing back to back. It was made the National Emblem of India in 1950.[246] The lion is also symbolic for the Sinhalese people, the term derived from the Sanskrit Sinhala, meaning "of lions"[247] while a sword-wielding lion is the central figure on the national flag of Sri Lanka.[248]

The lion is a common motif in Chinese art; it was first used in art during the late Spring and Autumn period (fifth or sixth century BC) and became more popular during the Han dynasty (206 BC – AD 220) when imperial guardian lions started to be placed in front of imperial palaces for protection. Because lions have never been native to China, early depictions were somewhat unrealistic; after the introduction of Buddhist art to China in the Tang dynasty after the sixth century AD, lions were usually depicted wingless with shorter, thicker bodies and curly manes.[249] The lion dance is a traditional dance in Chinese culture in which performers in lion costumes mimic a lion's movements, often with musical accompaniment from cymbals, drums and gongs. They are performed at Chinese New Year, the August Moon Festival and other celebratory occasions for good luck.[250]

Western world

Dorothy Gale meets the Cowardly Lion in The Wonderful Wizard of Oz. Art by W. W. Denslow, 1900.

Lion-headed figures and amulets were excavated in tombs in the Greek islands of Crete, Euboea, Rhodes, Paros and Chios. They are associated with the Egyptian deity Sekhmet and date to the early Iron Age between the 9th and 6th centuries BC.[251] The lion is featured in several of Aesop's fables, notably The Lion and the Mouse.[252] The Nemean lion was symbolic in ancient Greece and Rome, represented as the constellation and zodiac sign Leo, and described in mythology, where it was killed and worn by the hero Heracles,[253] symbolising victory over death.[254] Lancelot and Gawain were also heroes slaying lions in the Middle Ages. In some medieval stories, lions were portrayed as allies and companions.[255] "Lion" was the nickname of several medieval warrior-rulers with a reputation for bravery, such as Richard the Lionheart.[231]

Lions continue to appear in modern literature as characters including the messianic Aslan in the 1950 novel The Lion, the Witch and the Wardrobe and The Chronicles of Narnia series by C. S. Lewis,[256] and the comedic Cowardly Lion in L. Frank Baum's 1900 The Wonderful Wizard of Oz.[257] Lion symbolism was used from the advent of cinema; one of the most iconic and widely recognised lions is Leo, which has been the mascot for Metro-Goldwyn-Mayer studios since the 1920s.[258] The 1966 film Born Free features Elsa the lioness and is based on the book Born Free published in 1960.[259] The lion's role as king of the beasts has been used in the 1994 Disney animated feature film The Lion King.[260]

Lions are frequently depicted on coats of arms, like on the coat of arms of Finland,[261] either as a device on shields or as supporters, but the lioness is used much less frequently.[262] The heraldic lion is particularly common in British arms. It is traditionally depicted in a great variety of attitudes, although within French heraldry only lions rampant are considered to be lions; felined figures in any other position are instead referred to as leopards.[263]

See also

Notes

  1. ^ Populations of India are listed in Appendix I.

References

Citations

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  2. ^ a b c d e f Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P. & Nowell, K. (2017) [errata version of 2016 assessment]. "Panthera leo". IUCN Red List of Threatened Species. 2016: e.T15951A115130419. doi:10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en. Retrieved 15 January 2022.
  3. ^ a b Linnaeus, C. (1758). "Felis leo". Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. Tomus I (decima, reformata ed.). Holmiae: Laurentius Salvius. p. 41. (in Latin)
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Lion: Brief Summary

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The lion (Panthera leo) is a large cat of the genus Panthera native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adult male lions are larger than females and have a prominent mane. It is a social species, forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator; although some lions scavenge when opportunities occur and have been known to hunt humans, lions typically do not actively seek out and prey on humans.

The lion inhabits grasslands, savannahs and shrublands. It is usually more diurnal than other wild cats, but when persecuted, it adapts to being active at night and at twilight. During the Neolithic period, the lion ranged throughout Africa and Eurasia from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in western India. It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have declined by about 43% since the early 1990s. Lion populations are untenable outside designated protected areas. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes for concern.

One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire and have been a key species sought for exhibition in zoological gardens across the world since the late 18th century. Cultural depictions of lions were prominent in Ancient Egypt, and depictions have occurred in virtually all ancient and medieval cultures in the lion's historic and current range.

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