African lion, Panthera leo August 2005

Fact Summary Taxonomy and Nomenclature Distribution and Habitat Physical Characteristics Behavior and Ecology Diet and Feeding Reproduction and Development Genetics Diseases and Pathology Managed Care Population and Conservation Status Web Resources Bibliography

Describer (Date): Linnaeus (1758) Syst. Nat., 10th ed., 1:41  

Kingdom: Animalia
    Phylum: Chordata
        Class: Mammalia
             Order: Carnivora
                  Family: Felidae (Fischer de Waldheim 1817)
                        Subfamily: Acinonychinae (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 1854)
                                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 groups.
• 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 / African lion
      Panthera l. leo (Linnaeus 1758) North Africa / Barbary lion / extinct
      Panthera l. massaicus Masai Lion Uganda and Kenya / Masai lion
      Panthera 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 species.
• Wild cats dispersed everywhere except Australia, Antartica and Oceana.
• Divergence between Asian and African lions occurred about 200,000 years ago.

Distribution 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 Habitat: 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

Measurements

• 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 cm)
• Height
  • M: 4 ft (123 cm)
  • F: 3.5 ft (107 cm)
• Tail length
  • 2-3.3 ft (60-100 cm)
    

• Only cat with tail tuft (both male and female)
  • Tuft 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 (McBride 1977)
  • Reported only in vicinity of Kruger National Park and the Umfolozi Game Reserve in South Africa
  • Eyes and skin are pigmented (leucism, NOT albinism)

• Size: males almost 50% larger than females
• Mane: only found in males
• Development (Schaller 1972)
  • Noticible around 2 years of age
  • Development strongly influenced by testosterone
• Functions
  • Indicates fitness to potential mates
  • Possible protection of neck
  • Individual recognition   
• Disadvantage in hunting - easy for prey to spot maned males

• 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
  • Thicker mane

• Canine teeth approximately 60 mm (2.4 in) long

• 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 are hunted
• 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 control
• 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 cubs
• 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 success
• 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 15-1,000 kg 
• 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 mechanisms
• 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 hours+)
• 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 age

• 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 defensive

• 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 situations

• 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 immediately afterward.
• 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 distances.
• Top running speed 48-59 km/hr (Chassin, et.al., 1976)
• Not adept climbers. Usually climb to play or rest, scan for prey or escape.

• 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 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 giraffes
• Schaller found lions in savanna killed 88% of food, lions on plains scavenged 53%
• 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 one meal.
• 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

• 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 attended closely
• March - July strongest birth peaks February - April weakest

• 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

• 110 days (100-119days) 

• 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)

• 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,

• 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
• Weaning
  • Mothers have no milk by time cubs are 7 months
  • But suckling may continue from other lactating females for up to 12 months

• 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 (Sunquist)
• 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

• 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

• 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.

• Occasionally gored by prey they attack (horns and tusks of prey evolved as defensive fighting organs)
• CDV - Canine Distemper Virus - 1993 epidemic in the Serengeti affected more than 1/3 of the 3,000 lions in the area.
• FIV-Pie - Feline Immunodeficiency virus - chronic species-specific strain carried by free-ranging lions has not been connected with immune pathology or mortality (Troyer 2004)
• Parasitic and other diseases: (Bjork 2000) (Horak 1987) trichinosis, filariasis, sarcoptic mange, pentastomiasis, echinococcosis, taeniasis, hepatozoonosis, anthrax, babesiosis and rickets
• Gastric spiral bacteria and intramuscular sarchocysts (Kinsel 1998)
• Ixodid ticks

• Obesity, inactivity and stereotypy (implementation of random gorge and fast eating schedule)
• FIV - Feline Immunodeficiency Virus - could be associated with neorologic, haematologic, and urologic abnormalities, and increased aggression and other behavioral changes. Transmission is poorly understood (Wack 2000)
• CDV - Canine Distember Virus (Appel 1994) (Endo 2004)
• Degenerative spinal disorders (Kolmstetter 2000)
• Cerebellar herniation
• Feline asthma syndrome (Mukherjee 1999)
• Description of viral diseases of non-domestic felids and treatment at: www.felidtag.org 

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.

• 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 Asian-African hybrids.
• 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 and 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 defects)
• 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 www.felidtag.org 

• Original 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)

• 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 Central Africa
• 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 1996).
• 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

• Lion Research Center (University of Minnesota): lion facts, research, conservation activities
• African Lion Working Group (ALWG): facts, research, news
• Asiatic Lion Information Center (Wildlife Conservation Trust): information on Asiatic lion and India's Gir ecosystem
• ARKive: African lion facts, images; videos of behavior in wild
• Felid Taxon Advisory Group (AZA): Panthera leo. African lion fact sheet
• IUCN/SSC Cat Specialist Group: rich but slightly outdated information source for 36 living cat species. Visit their Cat Digital Library for downloadable documents relevant to conservation of wild cats (SDZG staff may obtain username/password from kelliott@sandiegozoo.org)
• IUCN Red List: Panthera leo. African lion fact sheet

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