African lion, Panthera leo
TAXONOMY & NOMENCLATURE
(O'Brien et al 1987)
(Shoemaker 1997) (Nowell & Jackson
Describer (Date): Linnaeus (1758) Syst. Nat., 10th ed., 1:41
Taxonomic History and Nomenclature
Family: Felidae (Fischer de Waldheim 1817)
(Pocock 1917) - cheetah
Subfamily: Felinae (Fischer de Waldheim 1817) - small and medium-sized non-pantherine cats
Subfamily: Pantherinae (Pocock
1917) - leopard, jaguar, lion, tiger, snow leopard
Genus: Neofelis (Gray
Genus: Panthera (Oken,1816)
Species: Panthera leo (Linnaeus 1758) - lion
Subspecies: Panthera leo leo - African lion
Subspecies: Panthera leo persicus (Meyer
1826) - Asian lion
Species: Panthera onca - jaguar
Species: Panthera pardus - leopard
Species: Panthera tigris - tiger
Genus: Pardofelis (Severtozov 1858) - marbled cat
Genus: Uncia (Gray 1854) - snow leopard
- 36 species of wild cat - reclassified many times with different groupings and generic names
- The taxonomy proposed by Wozencraft is used by CITES
and IUCN and lists Panthera leo as a monotypic species. (Combining the
Asian and African populations)
- O’Brien, et al, determined that African lions contain 3 unique
alleles not found in the Asiatic population. He determined that the
genetic difference between Asian and African lions is equivalent to the
distance between different subspecies of tiger, cheetahs and human racial
- The most recent mitochondrial study (Dubach) found modest genetic
variation among African lion populations but possibly 4 evolutionary
significant units (ESUs)
- ISIS - The International Species Inventory System, observes a
sub-species classification based on a proposal by Shoemaker in 1997:
Panthera l. bleyenberghei (Lonnberg 1914) Angola &
Zimbabwe / African lion
Panthera l. krugeri (Roberts 1929) South Africa /
Panthera l. leo (Linnaeus 1758) North Africa / Barbary
lion / extinct
Panthera l. massaicus Masai Lion Uganda and Kenya / Masai lion
l. melanochaitus (Smith 1858) Cape province / extinct
Panthera l. nubicus (Blainville
1843) Tanzania / East African lion
Panthera l. persicus Gir
Forest, India / Asian lion
Panthera l. senegalensis (Meyer
1826) Senegal - Cameroon / African lion
- Collective noun "pride" dates from mid-15 century
||From Haas, S. et al, 2005.
Geographic distribution of Panthera leo in Africa and India.
1. P. l. azandicus
2. P. l. bleyenberghi
3. P. l. krugeri and P.l. melanochaitus
4. P. l. nubicus
5. P. l. persica
6. P. l. senegalensis
- Common names: African lion; Asian lion
- Title: "The King of Beasts"
- From the lion's regal bearing and impressive mane
- The larger, more powerful
tiger was not known to Europeans when they first encountered lions
- Other European names:
- Lion d'Afrique (French); león (Spanish); Löwe (German)
- Some African names:
- Ingonyama (Xhosa, Zulu); simba (Swahili); shumba (Shona); sotho (Tau)
- The International Foundation for the Conservation of Wildlife lists 160
different names used by native Africans
- The Felidae and Canidae families diverged from a
common ancestor about 50 million years ago. Felid phylogeny has been
difficult to analyze because many species have split from each other
relatively recently: 10-15 million years ago.
- The Felidae karyotype is highly conserved. The
domestic cat has 19 chromosome pairs, 15 of which are the same in all cat
- Wild cats dispersed everywhere except Australia, Antartica and Oceana.
- Divergence between Asian and African lions occurred about 200,000 years
DISTRIBUTION & HABITAT
(Bauer & van der Merwe 2004) (Sunquist 2002) (Nowell &
- Historically found from Africa through Eurasia and
North America into South America
- Currently found only in sub-Saharan Africa (Botswana, Ethiopia, India,
Kenya, Malawi, Mali, Mozambique, Namibia, Senegal, Somalia, South Africa,
Sudan, Uganda, Zambia, and Zimbabwe) and the the protected Gir Forest in
northwestern India. The majority are in East and Southern Africa.
- Link to IUCN map
- Broad habitat tolerance - most altitudes and all
vegetation types except tropical rainforests and interior of Sahara
desert. Able to live on moisture from prey or plants
- Optimal habitat : open woodlands, and thick bush, scrub and grass
complexes, with some cover for hunting and denning.
- More at home in open areas than tigers, jaguars, or leopards
- Prey requirements: Medium- to large-sized ungulates (buffalo,
antelopes, zebra, and wildebeest)
From Bauer, H, and van der Merwe, S, 2004
(Grzimek 1990) (Eaton 1974) (Estes 1990) (Londei 2000) (Rudnai 1973) (Schaller 1972) (Sunquist 2002)
(Williams et al. 1997)
- Body weight
- M: 330-573
lbs (150-260 kg)
- F: 265-397 lbs (120-180 kg)
- Head and body length
- M: 5.6-8.2 ft (170-250 cm)
- F: 5.25-6.23 ft (160-190
- M: 4 ft (123 cm)
- F: 3.5 ft (107 cm)
- Tail length
- Only cat with tail tuft (both male and female)
covers horny spur at tip of tail (Rudnai 1973)
- Only male cat with mane (male tiger has a ruff)
- Eyes set laterally on head provide good wide-angle vision
- Ears rounded
- Inner ear has long mobile pinna able to localize sound source
- Large nostrils and complex nasal passages
- Massive limbs built more for attack than running
- 4 teats
- Coat: tawny
- Tail tuft, patches on the back of ears, and lips: black
- Cubs are spotted
- Color change begins around 3 months, but some spots may
persist to adulthood
- White pelage is unusual
- Reported only in vicinity of Kruger
National Park and the Umfolozi Game Reserve in South Africa
- Eyes and skin are pigmented (leucism, NOT albinism)
Differences between Asiatic and African lions
- Size: males almost 50% larger than females
- Mane: only found in males
- Noticible around 2 years of age
- Development strongly influenced by testosterone
- Indicates fitness to potential mates
- Possible protection of neck
- Individual recognition
- Disadvantage in hunting - easy for prey to spot maned males
Teeth (Sunquist, 2002)
- Asiatic lions
- Belly fold - flap of skin that runs the length of the
belly between the front and hind legs
- Thicker coat, longer tail tassel
- African lions
- Canine teeth approximately 60 mm (2.4 in) long
BEHAVIOR & ECOLOGY
1972) (Mellen 1993) ( Estes 1991) (Kingdon 1977) (Packer & Pusey 1997) (Nowell &
- Least active of all felids
- Hunt, feed and mate at all times. Behavior is influenced by season,
habitat, and humans
- Nocturnal in areas where there is little cover to stalk prey or they
- 20-21 hours spent resting (sleeping difficult to ascertain in the wild.
Lions in zoos sleep 10-15 hours) (Haas 1958)
- 2 hours walking (average = 9.5 km/day)
- Daily movement erratic and unpredictable. Remain in
an area as long as there is prey
- Activity of cubs, subadults and males is
based on movement of adult lionesses
- Extremely tolerant of sun. Move in and out of shade for temperature
- 5 pm - 7 pm: (dusk): stretch, defecate, groom, play, rub cheeks
- 7 pm - 6 am: roaming, eating (peaks at dusk and dawn) resting
- 8 am -5 pm: resting, shift resting sites from sun to shade
- Extremely social (tigers and leopards are solitary);
cheetahs are the only other non-solitary felid (Mellen)
- Coordinated group possibly more successful in killing large prey
- Better defense / raising of young
- Maintenance of territory
- Insurance against individual injury or temporary incapacity
- Pride is social unit = Matriarchal society of related females and their
- Average size: 4-6 adult females. Smaller in arid environments.
- (Pride of 20 females observed where prey was abundant year-round)
- No evidence of hierarchy
- Membership is stable but individual members often
scatter in small foraging sub- groups throughout the pride's range.
Individuals may spend considerable time alone. A
"fission-fusion" social unit.
- Pride members often give birth synchronously; young are raised
communally in what is termed a "creche"
- Males (up to 8) form lifelong alliances to maintain breeding rights
over one or more prides.
- Males remain aloof from group interactions except
where food is concerned
- Adult males leave spontaneously, (few months - 3 years) or are ejected
by new males.
- Visual hunters - wind direction is not a factor in
- Females do most of the hunting. (Males less agile - can't run as fast)
- Hunt mainly at night. Most prey consumed immediately.
- Preferred Prey: mammals 50-300kg. Will attack any animal between
- Large animals are strangled, smaller ones
bitten deeply in head neck or chest.
- Medium-sized prey grabbed by rump
with forepaws and brought down. Killing bite to throat strangles prey (may hold for as
long as 10 minutes). While one lion grasps throat others begin to eat.
- Small prey felled with
slap to thigh or grabbed with both paws. Immediately bitten. Killing bite
to back of neck, head, throat.
- Pattern: stalk, then quickly charge.
- Schaller's observation on Serengeti lions: 48% single hunter, 20% 2
hunters, 32% communal hunts.
- Cubs join in hunts by 15 months. Become fully-skilled at 2.5 years
- Sometimes kill more prey than they can consume. Stay near kill to feed
later. May move small prey to cover to eat protectedly.
- Each lioness in a given pride repeatedly occupied the same position
in hunting formation during group hunts (Stander).
- Both sexes scent-mark
- Spraying and marking define territories - function as spacing
- Patrolling and roaring used to maintain territory
- Home range of a pride is largely set by biomass of prey available:
- From 20 sq km to 200 sq km
- Friction among pride members is rare. Fights are
short and biting infrequent - mainly vocalization and teeth-baring.
- "Kills" are disruptive to pride harmony.
- Males will
appropriate a small kill from a female.
- Lionesses commonly take meat from cubs.
- Although smaller than males, females can react aggressively toward
them. Several females may gang up to route a male
- Infanticide can occur when a new coalition of males takes over a pride.
- More than 1/4 of all cubs are killed by invading males (Sunquist).
- Lionesses may be killed trying to defend their cubs.
- Cubs usually left unguarded (24
- Mothers suckle, guard and transport cubs for first few weeks.
- Communal nursing by all lactating lionesses
- Cubs subsist entirely on milk for first two months - gradually
supplemented wtih meat beginning around 5 weeks
- Males often prevent cubs starvation: Will control remains of a carcass
preventing lionesses from eating but allowing cubs to feed. (Cubs usually
arrive late to kills)
- Males taking over a pride will attempt to kill any cubs under 1 year of
- Head-rubbing - a friendly greeting is common (rare in cheetahs)
- Social licking common between females and cubs
- Head-low posture denotes threat
- Male strutting denotes dominance / display
- Ears flattened, teeth bared, eyes narrowed crouching, lying on back is
- Used for territoriality, intimidation, social bonding, advertising presence
- Only lions and tigers can roar. Snow leopards and cheetahs do not roar.
- Mechanism: Hyoid apparatus is incompletley ossified. The attached elastic
ligament can be stretched to create the roar (Hast 1989)
- Males begin roaring at 1 year - females a few months later
- Gradations in volume - Full roar is audible for up to 8 km
- Lions can roar from any position but usually standing
- Other sounds: growl, snarl, hiss, meow, grunt, and puff.
- Puffing, which sounds like a stifled sneeze, is used in friendly
- Rubbing: Rub muzzle
on vegetation (tuft of grass, shrub, sapling etc.) prior to marking. Males
pick up odors and later pass them to other pride members
- Spraying : Caudally directed penis sprays urine (mixed with scent
from 2 anal glands near the base of tail). 1-20 squirts / marking (Female
lions rarely spray unlike female tigers) Commences around age 3.
Fifty-five compounds found in urine, including 7 potential species
identifying compounds (Andersen 1999)
- Scraping: Claws of hind paws alternately rake the ground from 2-20
times. Paws leave a concentration of scent. Urination occurs during or
- Common in both male and female beginning around age 2.
- Frequency depends on: vegetation in habitat, males responding to
estrous lionesses, vicinity of kills
- Feces: Deposited randomly. Probably not part of marking system like
tiger and puma
- Walking speed: 3-4 km/hr
- Little stamina - unable to pursue an animal rapidly over long
- Top running speed 48-59 km/hr (Chassin, et.al., 1976)
- Not adept climbers. Usually climb to play or rest, scan for prey or
- Interact with hyenas, jackals and vultures who
compete for remains of a kill
- Readily appropriate kills made by hyenas
- Generally ignore jackals but will attack hyenas
DIET & FEEDING
(Schaller 1972) ( Estes 1991) (Sunquist 2002)
- Diet found to include 38 different species
(Influenced by prey abundance - will eat whatever is more available) Also
effective scavengers - forcing other animals off "kills"
- Diet ranges from rats, reptiles, fish, groundnuts to rhinos and bull
- Schaller found lions in savanna killed 88% of food, lions on plains
- Main Food Items eaten by lions in 4 African Reserves between 1954
-1969: buffalo, wildebeest, zebra, impala, waterbuck, hartebeest, warthog
(can dig them out of burrows), giraffe. Gir lions feed mainly on chital
deer - occasionally sanbar, nilgai and domestic stock.
- All parts of body consumed except for contents of digestive tract,
horns, teeth and occasionally lungs. If a group is unable to eat all the
meat at once, 1 or 2 (usually males) remain to guard the kill
- Males can eat as much as 40 kg or one-quarter of their body weight, in
- Drink daily if water is available but often go 4-5 days without water
- Composition of milk is detailed by De Waal 2004
- Diet information for zoo animals is described by Crissey 2003
REPRODUCTION & DEVELOPMENT
(Packer & Pusey, 1984 )(Schaller 1972) (Estes 1991)
(Nowell & Jackson 1996)
- Females are polyestrous (cycle throughout the year)
Like most cats they are induced ovulators.
- Fights over a lioness in heat by male members of a pride are uncommon.
- Copulation initiated by male or female
- Female in estrous exhibits extreme restlessness: lying down, jumping
up, walking rapidly, rolling and twisting on her back.
- Neck-bite common in many mating cats (probably to
induce temporary passiveness in female) has become ritualized in the lion.
(Light, brief bites in 60% of observed matings)
- Contact terminates quickly after copulation (copulation lasts 8-68
seconds, averaging 21 secs. and occurs about once every 25 minutes.)
- Estrous females may accept several males in succession if she is not
- March - July strongest birth peaks February - April weakest
Estrus: 4 days
- Average tenure of male coalitions in prides is only
24 months. Following a pride takeover, the incoming males kill small cubs
and chase out young inducing females to come into estrus within a few weeks
- Females conceive an average on 138 days after the loss of cubs, whereas
females with surviving young will not mate again until their young are more than 18 months old
- Females will defend their cubs
Interestrus Interval: 16 days
- Birth occurs in a den or dense brush.
- Weight: 1.2-2.1 kg.
- Eyes open at birth or shortly after
- Fur is spotted (spots on belly and legs)
Litter size: 1 to 4 cubs
- 20 months (11-25 months) if previous litter survives
- 4-6 months if litter is lost. (Pusey & Packer, 1987)
- Births are synchronized by male take-overs
- Sex Ratio: At least 2:1 favoring females. Possibly an adaptation -
males don't hunt, eat more than their share and a large number of
males are not needed in a polygynous society.
- More than half the cubs don't survive the first year,
Large Cubs 1-2 years
- Walking by 10 days
- Remain hidden until mobile - around 4-6 weeks
- Mother and cubs rejoin pride around 6-8 weeks - cubs raised communally
- First set of milk teeth erupt around 3 weeks, canines at 4 weeks
- Milk is main diet until solid food at around 2 months,
- Cubs will suckle from any lactating female...but females try to give milk
primarily to their own young
- Eyes change from gray-blue to amber at 2-3 months
- 3-5 months acquire adult coat
- Tail tuft, absent at birth develops between 5-7 months
- Males may have slight throat ruff at 6 months, stockier than females
- Starvation / predation / infanticide / abandonment common causes of death
- Mothers have no milk by time cubs are 7 months
- But suckling may
continue from other lactating females for up to 12 months
Subadults 2-4 years
- Cubs are leopard size - 45 kg
- Deciduous canines replaced with permanent canines at 13.5-15 months
- Mane in males consists of crest on head and nape with tufts on cheeks
- By the time they are 2 years of age lionesses are competent hunters
- Females slender, with short muzzle, inconspicuous nipples, taut abdomen
- Males undergo a 6 month growth spurt around 3.5 years, with light blond
beginning of mane
- Males leave birth pride (social group) around 3.5 years
Young Adults 4-6 years
- Both males and females reach sexual maturity around 24 months
- Males may not produce sperm until 30 months. In the wild, they may not
have an opportunity to reproduce until the age of 5
- Most females have first litter around age 4
- Most males leave the pride by the age of 4
- Mane clearly present. Brownish with
rust, yellow and black hairs
- Both males and females continue to grow until age 6
- Females can breed until they are 15 years old but reproduction begins to
decline at about 11 years
- Males as old as 16 can produce viable sperm, but reproduction probably ceases
after pride tenure is lost
- In the wild: Males live 12-13 years, females up to 18 years
- In captivity: 20-25 years (both sexes)
- Lioness at Cologne Zoo lived to age 30
(Guggisberg 1975) (O'Brien, 1987) (Dubach, 2005)
- 38 chromosomes - 19 pairs
- Have been crossed with tigers, leopards, jaguars (liger if father is a
lion; tigon if father is a tiger)
- Asiatic and African lions are genetically distinct although the
difference is smaller than
the genetic distance between human racial groups.
- Phylogenetic data from a 2005 study found 6 distinct haplotypes forming
2 clades (eastern and western savanas)
- At least 4 lion groups
- the southwestern populations
- the populations
to the east of the rift valley
- the populations to the west of the rift valley
- the Sabi Sands population.
These regions might define evolutionary significant units (ESU) but modest
genetic variation argues against taxonomic distinctions.
(Livingston1974) (Kisling 2000) (Shoemaker www.felidtag.org) (CatNews 2004) (Green, R. 1991)
Imported by the Romans from North Africa for menageries
and public games. Thousands slaughtered for public entertainment. With the decline of Roman Empire, menageries remained in the hands of royalty
and the newly powerful catholic church. In the middle ages, public European menageries with lions began to spring up.
ISIS captive population
- 1068: First permanent English managerie belonged to Henry I
- 1235: Henry III kept and bred lions in the Tower of London
- 1716: A lion was first exotic animal exhibited in North America - Boston, MA.
Housed at the home of Captain Arthur Savage, later moved and exhibited at the
home of Martha Adams (1720) before touring major New England cities
- 1791: 2nd lion imported to US
- 1874: Philadelphia Zoo opened with a lion exhibit
- 1916: The roar of a lion from the Panama-California Exposition in San Diego,
inspired Dr. Harry Wegeforth to establish the San Diego Zoo.
- 1977: International studbook for Asian lions begun.
- 1983: Species Survival Plan (SSP) for Asian lions
- 1986: Discovery that all Asian lions in the U.S., except one, are
- A new breeding program is initiated using lions whose lineage can be
accurately traced back to the Gir Forest in India.
- 1992: African lions added to Regional studbook and added to SSP (Krugeri
nubicus are the only African sub-species identified in North America and Krugeri
is the only sub-species with long-term breeding potential if no others are imported)
- 2004: Central Zoo Authority of India orders more than 300 hybrid lions held
in zoos and safari parks across India sterilized (hybrids suffer from mental and physical
- 2004: "Lion Camp" opens at The San Diego Wild Animal Park featuring
a 6-member pride of lions. The 33,000 sq ft exhibit has a 40-foot-long glass viewing
window Regular demonstrations of current animal training techniques
- Zoo Standards for keeping large felids in captivity can be found at
POPULATION AND CONSERVATION STATUS
(Nowell and Jackson 1996) (Cat News 2004/2005) (Bauer & Van Der Merwe 2004)
range: large parts of Africa,
Europe, Middle East and Asia
- Disappeared from eastern Europe almost 2,000 years ago (A.D. 100)
- Disappeared from North Africa, Middle East and Asia between 1800 -1950
- 250 mature adults survive in India’s Gir Forest Reserve
- 22,600 remain in Africa (10% of the number present 25 years ago)
- West and Central Africa: small, isolated declining populations
- East and Southern Africa: many large populations stable over last 3 decades
- Serengeti: recovered from 1994 distemper epidemic (30% mortality)
Threats to survival
- IUCN Status:
- African lion = Vulnerable. Species population reductin of 30%-50% over
the past 2 decades (3 lion generations). Recent surveys have shown that
the lions of West Africa are in serious decline - the number of mature
individuals have been estimated by two separate recent surveys at 850
(Bauer and Van der Merwe 2004) and 1163 (Chardonnet 2002). Both
estimates are well below the Endangered level of 2500.
- Asian lion = Critically Endangered. Single population of ~250 mature
breeding individuals, all occurring within one area. Numbers are expected
to suffer a decrease due to increasing conflict with people.
- CITES Status:
- African lion: Appendix II
- Asian lion: Appendix I
- Habitat loss/degradation
- Isolation of population of West Africa population from populations of
- Conflict with human settlement/agriculture/cattle raising
- Lion predation on livestock
- Scavenging behavior makes them vulnerable to poisoned carcasses
- Many people are killed by lions resulting in retaliation
- Hunted for meat
- Migratory ungulate prey base creates lean seasons (Nowell and Jackson
- Ongoing war and civil unrest in Africa
- Disease from domestic dogs
- In Sub-Saharan Africa, the lion conservation community works in the
context of four regions : West, Central, East, and Southern.
- Management Plans developed by IUCN Cat Specialist Group and AZA
Felid Taxon Advisory Group :
- Habitat corridors needed
- Monitoring of protected areas
- Community education
- Captive breeding / Artificial propagation
- "Project Life Lion" is a major campaign to vaccinate as many
dogs as possible to guard against canine distemper.(Sunquist)
- Hunting is banned in Angola, Cameroon, Congo, Gabon, Ghana, Malawi,
Mauritania, Niger, Nigeria, and Rwanda. Hunting is restricted to
"problem" animals over in Benin, Botswana, Burkina Faso, Central
African Republic, Ethiopia, Ivory Coast, Kenya, Mali, Mozambique, Senegal,
Somalia, Sudan, Tanzania, Togo, Uganda, Zaire,
Zambia and Zimbabwe (Nowell & Jackson 1996).
- Gir Population: National Protection by Indian Government
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