African and Asian Lions (Panthera leo) Fact Sheet: Behavior & Ecology

Daily activity patterns (Schaller 1972; Rudnai 1973; Paul Funston, personal communication, 2016)

• Hunt, feed, and mate at all times of day
• Hunt actively at night, searching for prey
• Hunt opportunistically during the day, often close to waterholes or river banks
• Peak activity after 5pm and before 8am
• Rest most of the day
• Some morning activity
• Groom and play around dawn and especially in the evening
• Typically rest in the shade but will tolerate sun if stalking
• Cool themselves by panting or lying on their backs, exposing white underbelly fur
• During the day, tend to remain in an area if prey are available
• Forage at night
• Cubs, subadults, and sometimes adult males follow movements of females

Behaviors to conserve energy

• May rest up to 20-21 hours per day, on average (Schaller 1972)
• Lions in zoos sleep 10-15 hours (Haas 1958)
• Active early and late in the day, and at night to avoid the heat of the day (Wilson and Mittermeier 2009; Estes 2012)
• Sleep and rest in shade, commonly in compact groups (Wilson and Mittermeier 2009)
• Interrupted by periods of intense activity (hunting, showing aggression)
• Rest for long periods when satiated (Wilson and Mittermeier 2009)
• Climb trees and lay along branches (Schaller 1972; Wilson and Mittermeier 2009)
• Cool themselves in a breeze
• Avoid flies
• Avoid dangerous animals
• Look around from a height

Often hunt at night (Wilson and Mittermeier 2009, except as noted)

• Conditions for easier hunting
• Little or no moonlight
• Stormy conditions, when noise, wind, and waving vegetation make it difficult for prey to detect lions
• Males may hunt as well as females, depending on location

Space use

• Mostly resident (Wilson and Mittermeier 2009)
• May follow migratory prey, seasonally (Wilson and Mittermeier 2009)
• Non-territorial males and subadults nomadic (Wilson and Mittermeier 2009)
• Varies by location
• May move a mean of 4.5-15 km in 24 hours, depending on location (Wilson and Mittermeier 2009)
• Males generally travel farther than females each day (Wilson and Mittermeier 2009)
• Females generally stay within natal area or an adjacent home range; do not disperse as far as males (Dolrenry et al. 2014)
• May disperse if their fathers are still resident when they mature (West and Packer 2013)
• Males taking over pride may also oust subadult females and males (Hanby and Bygott 1987; West and Packer 2013)
• Males disperse from their natal area; travel 2-3 times farther than females (Dolrenry et al. 2014, except as noted)
• Exception: Kruger National Park (Funston et al. 2003)
• 80% of young male coalitions remained close to natal territories
• Little known about lion dispersal in human-dominated areas (Dolrenry et al. 2014)
• Tend to select wooded areas with low lion densities (see publications by Nicholas B. Elliot and colleagues)

Home ranges

• May overlap or be exclusive (i.e., defended, a “territory”) (Wilson and Mittermeier 2009)
• Central area defended for exclusive use (Mosser and Packer 2009; West and Packer 2013)
• Home range size influenced by prey abundance, prey movements, and geography (Wilson and Mittermeier 2009; West and Packer 2013)
• Larger home ranges when less prey available
• Smaller home ranges when more prey available
• Home range size varies from 25-1000+ km² (average: about 100-200 km²) (Wilson and Mittermeier 2009)
• May be much larger in areas of low prey density
• Etosha National Park: up to 2075 km² (Stander 1991)
• Kgalagadi Transfrontier Park: 266-4532 km² (Funston et al. 2001)

Territorial Behavior

• Both sexes defend the territory (West and Packer 2013, and as noted)
  • Males range throughout the territory
    • Concentrate on edge areas and overlap zones
  • Females defend a core area against other lions
  • Larger prides more effectively gain disputed territory (Mosser and Packer 2009)
• Roaring (Wilson and Mittermeier 2009)
  • May be heard as far as 5 km (3 miles) away, under certain conditions
    • Advertises that a territory is occupied and holders are willing to defend it against intruders
    • May be individually distinctive, particularly when adult males are with the lionesses
  • Prides may roar as a group
• Patrol territory (West and Packer 2013)
  • Males patrol the extent of the territory, especially edge areas
  • May walk beyond their ranges, but usually retreat at detection of resident lions
  • Avoid fights
    • Use caution when outnumbered
• Scent marking
  • Leave urine and scat, and scrape the ground to mark territory (Wilson and Mittermeier 2009)
  • Scent marks concentrated along territory edges and in areas of higher probability of pride overlap (e.g., near waterholes or rivers) (Paul Funston, personal communication, 2016)
  • Urine marking mostly performed by males (Estes 2012)
    • Females occasionally spray

Prides as social units

• Only truly social cats (Mosser and Packer 2009; Wilson and Mittermeier 2009, except as noted)
• Other cat species (e.g., tigers, leopards) are solitary, except cheetah (Mellen 1993)
• Fission-fusion social dynamics (Mosser and Packer 2009; West and Packer 2013)
• Dynamic social interactions; appear to have preferential associations (Abell et al. 2013)
• Individuals found in a number of subgroups (Mosser and Packer 2009)
• Cohesive social bonds among members (Abell et al. 2013)
• Often based on kinship
• ‘Keystone individuals’ may be important in maintaining group cohesion
• Lions form matriarchal groups called “prides” (Wilson and Mittermeier 2009, except as noted)
• Include genetically related females and their cubs
• No known dominance hierarchy among females
• Often give birth synchronously
• Communally raise young in what is termed a crèche
• Nurse one another’s cubs (“allosuckling”)
• Most common among closely related females or females with only one cub (Pusey and Packer 1994)
• May improve chances of protecting offspring from intruder males and neighboring prides (West and Packer 2013)
• Cooperative hunting
• Proposed benefits of sociality (Wilson and Mittermeier 2009, and as noted)
• Maintain and compete for territories (Mosser and Packer 2009)
• Cooperative defense of the pride and of cubs
• Help in cub rearing
• Defense of kills
• Improved success in hunting large prey
• Less affected by injury, death, or incapacitation of individuals
• Average pride size
• 3-10 adult females, plus their offspring; 2-3 resident males (West and Packer 2013)
• Larger groups have a competitive advantage (Mosser and Packer 2009)
• Larger in productive areas
• Serengeti National Park: 1-21 adult females, plus their offspring; 1-9 adult males (Mosser and Packer 2009; West and Packer 2013)
• More intense competition for territory (Mosser and Packer 2009)
• Smaller in very arid environments (~2-3 adult females)
• P. l. persica: 2-5 adult females, on average (Nowell and Jackson 1996; Wilson and Mittermeier 2009)
• Pride membership is stable (Wilson and Mittermeier 2009)
• Individual members often forage in smaller subgroups throughout the pride's range
• Known as a "fusion-fission" social system
• Members may spend considerable time alone

Male membership

• A single male or up to 7 males may join a pride (Wilson and Mittermeier 2009)
• Young males raised from the same natal crèche may form lifelong alliances (West and Packer 2013).
• Groups called “coalitions”
• Cooperatively maintain breeding rights over one or more prides
• Young males may also be alone due to lack of male crèche mates or death of companions (West and Packer 2013)
• Single males may form a coalition with another single, unrelated male
•  Behaviorally indistinguishable from groups of related males
• Male coalitions challenge each other to hold residency with a pride (“tenure”) (Mosser and Packer 2009)
• Winners hold territory and mating rights
• Exclude strange males from siring cubs
• Larger groups can ‘hold tenure’ more than twice as long as a single male or pair of males (Wilson and Mittermeier 2009)
• Larger groups usually related
• Pairs sometimes unrelated
• Remain with pride, on average, only 2-3 years; may be less (Wilson and Mittermeier 2009, except as noted)
• Tenure duration generally increases with coalition size (West and Packer 2013)
• May be ejected by new males or leave in search of a new pride
• Large coalitions often move to adjacent pride areas  (West and Packer 2013)
• May defend more than one prides' territory (Wilson and Mittermeier 2009; Paul Funston, personal communication, 2016)
• Males that leave may become nomadic (West and Packer 2013)
• Hold pride tenure and sire offspring for only about 6 years during lifetime (Funston et al. 2003)

Other social aspects

• Males tend to remain aloof from group interactions, except where food is concerned (Schaller 1972)
• Males are successful hunters, often doing their own hunting (Funston et al. 1998)

Infanticide

• Common if new, intruder males take over a pride (Wilson and Mittermeier 2009)
• Assures males of their paternity (Wilson and Mittermeier 2009)
• Disadvantageous for females (Packer and Pusey 1983)
• Females defend cubs, often unsuccessfully (Packer and Pusey 1983; Wilson and Mittermeier 2009)
  • May gang up on intruder males (Estes 2012)
  • May be wounded (Packer and Pusey 1983)
  • Sometimes killed while caring for their cubs (Mosser and Packer 2009)
  • Many lose cubs within a month of takeover (Wilson and Mittermeier 2009)
    • In Serengeti: more than 1/4 of all cubs are killed by invading males (Sunquist and Sunquist 2002)
      • Infanticide less frequent in other areas
  • Pregnant females lose cubs shortly after giving birth (Packer and Pusey 1983; Wilson and Mittermeier 2009)
• Following infanticide
  • Period of adult female infertility followed by heightened sexual activity (Packer and Pusey 1983)
    • Occurs in first few months after takeover
• Also see Reproduction
• Removal of males via trophy hunting may potentially exacerbate infanticide rates, and thus, reduce population size (Packer et al. 2009)

Avoiding conflict (Estes 2012)

• Olfactory cues and vocalizations enable nomads and trespassers to avoid encounters with territory holders
• More effort invested in intimidating, deterring, and driving away strangers than fighting
  • Strangers may be escorted away from territory
• Biting generally avoided unless needed
  • Show little restraint when biting
  • Curbs potential injuries
     

Infrequent aggressive acts

• Live amicably, for the most part (Schaller 1972)
• Fights generally short (West et al. 2006)
  • Non-lethal vocalizations and teeth-baring prevent escalation (Schaller 1972)
  • Injury typically avoided
• Fight behaviors: slapping, grappling, biting, group attacks (Estes 2012)
  • Forehead and hindquarters often targets (West et al. 2006)
• Fights between pride members (Schaller 1972; Estes 2012)
  • When competing for meat
    • Increased aggression at kill sites
    •  “Share food, but often grudgingly”
    • Weaker lions must persist in gaining their share of meat
    • Males may monopolize smaller kills, taking them from females
      • Females, though smaller, will react aggressively
  • When competing over females in estrus
    • Female may provoke by seeking attention of a second male
• Fights between coalition males (West et al. 2006)
  • Swat and hit each other on the face
  • Rarely goes beyond slapping
• Fights with strange males (West et al. 2006, except as noted)
  • Assess when outnumbered
    • Avoid fights when low odds of winning
  • Attempt to incapacitate rivals
  • Generally only fight if immediate threat
    • Fight without hesitation if territory, females, or other valuable resource are threatened
  • Fights often severe and fatal (Estes 2012)
    • Death often quick for at least one participant 

Conditions necessary for play (Schaller 1972)

• Temperature not too hot
• Absence of hunger
• Secure environment or protection of adults
• Cubs feel safe with lionesses present

Some types of play (Schaller 1972)

• Chasing
• Wrestling
• Pawing
• Stalking
• Rushing

Cubs

• More playful than subadults and adults (Schaller 1972)
• Attempt to engage any adult (Schaller 1972; West and Packer 2013)
• Climb on females
• Jump against adults’ sides
• Grapple with adults’ legs
• Chase adults’ tails
• Adults may ignore cubs, repel with snarls, or move away when annoyed
• Reprimand with a sharp growl
• Wrestle frequently from 2-6 months old (Schaller 1972)
• Transition to more chasing and stalking between 6 and 12 months old
• Wrestling becomes less frequent with age
• Rare in adulthood
• Will play alone (Schaller 1972)
• Also, with objects
• Sticks, bark, tufts of grass, feces, bones
• Practice hunting skills
• Stalk, pounce, grapple, and chase in play (Wilson and Mittermeier 2009; Estes 2012)
• Develops motor patterns used in hunting
• Learn to hunt by watching adults (Estes 2012)
• Ambushing skills
• Observe prey movements and when adults ambush prey
• Begin participating in hunts around 11 months old
• Young cubs on hunts may spoil ambush attempt for adults (Schaller 1972; Estes 2012)
• Alert prey by running ahead, vocalizing, or playing
• Frequency of play behavior declines after a year of age (Rudnai 1973)

Adults

• Females never lose playfulness, but males older than 3 years rarely join in (Rudnai 1973; Estes 2012)
• Females engage cubs in play (e.g., pawing, chasing) (Rudnai 1973)
• Play with adult males infrequent
• Clawing, climbing, pawing, mouthing, and rolling behaviors observed with prey carcasses (Schaller 1972)

Tactile communication

• “Greeting ceremony” (Rudnai 1973; Estes 2012)
  • Approach each other, sometimes with soft moans; then rub heads and sides of body together
    • Tails raised or draped over the other lion’s back
    • Lean against each other
  • May demonstrate pride membership, reinforce social bonds, or be an effort of subordinates to appease more dominant individuals
  • Greetings between females and by cubs-to-mothers are common
  • Cubs and females may initiate greetings with males
    • Males usually reserve greetings for other males
• Head rubbing
  • Thought to maintain and strengthen social bonds (Matoba et al. 2013)
    • Avoided between partners after fights or tense situations (Schaller 1972)
  • Commonly observed in the wild (Schaller 1972)
  • Frequency varies between genders
    • Most frequently observed between two males, in one managed care study population; indicates strong social bonds in male coalitions (Matoba et al. 2013)
    • Second most frequently observed: females towards males
    • Often reciprocal between females, but not usually males
• Social licking (Estes 2012, except as noted)
  • Less prevalent than head rubbing
  • Any pride member can initiate a licking session
    • May be carried out by one or both partners
  • Cubs licked all over
  • In adults, licking typically on the head, neck, and shoulders
  • More commonly observed
    • Between mothers and cubs
    • Between two females, in one managed care study population (Matoba et al. 2013)
      • Seems to support function of social bonding (rather than expressing social status or reducing tension between lions)
  • Olfactory aspects of tactile communication need further research (Matoba et al. 2013)

Visual communication (Estes 2012)

• Use facial expressions, eyes, ears, tail, and body postures to convey mood and behavioral intent
  • Often combined with vocalizations
  • Associated with flight, defense, and attack
  • Facial expressions may or may not synchronize with body postures
    • Mood registers in the face first
  • See Schaller (1972), p. 92-98 for descriptions
• Black lips, tail tip, and markings on backs of ears make expressions more noticeable
• Facial expressions and ear movements similar to other cats
• Displays
  • Offensive/threat
    • Head-low
      • Head low to ground
      • Shoulders higher than normal
      • Forelegs apart
      • Gazes at opponent
      • Tail whipped up and down
    • Mouth agape
      • Mouth slightly open
      • Lips pulled back straight
      • Eyes wide and pupils small
      • Ears twisted so black marks are visible to opponent
    • Male strut
      • Dominance display to females
      • Performed by adult males and cubs during play
      • Tall, stiff posture while walking and presenting his side to female; tail lifted over his back
  • Defensive
    • Ears flattened
    • Teeth bared
    • Eyes narrowed
    • Crouching or lying on back

Vocalization

• Communicate information or emotional state by changing volume, intensity, tempo, and tone of calls (Estes 2012)
  • E.g., a cub producing light meows to signal distress
• Roaring (Wilson and Mittermeier 2009)
  • Sequence
    • Begins with a few subdued, spaced-out calls
    • Then, calls of longer duration
      • Accelerated pace and increasing volume
    • Ends with shorter calls
  • Young cubs may join in the “chorus” of their pride with little mews
  • Other felid species do not regularly join one another in intense calling
• Vocal communication in cubs
  • Have similar vocal repertoire to adults, though modified (Schaller 1972)
  • Kittens can hiss and spit shortly after birth (Wilson and Mittermeier 2009)
    • Also purr
    • Little change in vocal qualities through development
  • React to mother’s calls up to 8-10 months in age (Rudnai 1973)
  • Ability to roar developed in adulthood (Wilson and Mittermeier 2009)
  • Lose ability to emit certain distress calls as adults (Wilson and Mittermeier 2009)
• Range of adult vocalizations (Rudnai 1973; Estes 2012, except as noted)
  • See Rudnai, Table V-5, for context and responses to vocalizations
  • Roar
    • Loud and booming; volume variable
    • Typically roar while standing
      • Can produce in any body position
    • Begins with a few moans
    • Followed by 4-18 thunderous roars
    • Ends with a series of grunts
    • Roar more frequently when active (night and dawn)
    • Respond to other lions’ roars
      • Also roar spontaneously
    • Possible functions
      • Advertises territory ownership or pride presence
      • Intimidates rivals
      • Strengthens social bonds (as when roaring in a chorus)
  • Grunt
    • Warning
    • Reinforce social bonds
      • Used by mother lions to call cubs
      • Contact call between nearby adults
  • Growl
    • Warning, threat
    • Cough: a short, explosive growl
  • Snarl and hiss
    • Emitted with mouth open
    • May transition into a spit
      • E.g., when a strange lion approaches
  • Meow, miaow (Wilson and Mittermeier 2009)
    • Helps keep contact at close range
    • Intense meow signals distress
  • Purr and hum
    • Sounds of contentment
    • Heard when cubs suckle and sometimes during copulation
  • Puff
    • Emitted through closed lips
    • Repeated as lions approach each other
    • Signals non-aggression
  • Woof
    • Made by startled lions
    • Expresses alarm

Scent Marking and Olfaction

• Important channel of communication with other lions (Wilson and Mittermeier 2009)
  • Smell one another’s groin region upon greeting (Estes 2012)
  • Chemicals persist on surfaces (Wilson and Mittermeier 2009)
    • Last for days or even weeks
  • Detectable at night and in dense cover (Wilson and Mittermeier 2009)
• Functions (Wilson and Mittermeier 2009)
  • Seems to be an assertion of rights to be at a place
    • Not ownership, per se: simply “area is occupied”
  • Separates individuals
    • Mark territory boundaries
    • Prevent direct contact
  • Bring individuals together
    • Mating
    • Guarding estrous females
  • Identify individuals
    • Exact information transmitted unclear
    • Possibly identity, sex, status, reproductive state, when last present in area, etc.
      • Urinary estrogrens indicate changes in a female’s reproductive state
    • May provide territory intruders with a means to assess the territory holder
  • Identify a group
    • Group members may share a common odor
  • No known odors that convey dominance
    • Dominant animals may mark more often or leave more scent
• Chemical odors found in lion’s saliva, urine, and feces, and scent glands (Wilson and Mittermeier 2009)
  • Little known about the chemistry of social odors in carnivores (Andersen and Vulpius 1999; Wilson and Mittermeier 2009)
  • Can smell scent traces left by another lion’s feet (Estes 2012)
• Ways to leave odor behind
  • Rubbing
    • Rub chins or bump foreheads in greeting (Wilson and Mittermeier 2009)
    • Rub head, cheek, neck, and body on objects or one another (Wilson and Mittermeier 2009)
    • Rub muzzle on vegetation (tuft of grass, shrub, sapling etc.) prior to marking (Schaller 1972)
    • Males might pick up odors and later pass them to other pride members (Schaller 1972)
  • Depositing urine and feces
    • Leave scent at conspicuous or elevated locations in home range (Wilson and Mittermeier 2009)
      • Strategically placed
      • Often at nose level
    • Commonly spray and/or scrape in the vicinity of (Schaller 1972):
      • Kills
      • Estrous females
      • Scavengers, such as hyenas
      • Rival lions
    • Spraying
      • Mainly performed by males (Schaller 1972; Estes 2012)
      • Seems to be preferred method of marking (Wilson and Mittermeier 2009)
        • Continually updated to communicate occupancy
      • Use backward-facing penis (Schaller 1972; West and Packer 2013)
      • 1-20 squirts per marking (Schaller 1972)
      • Urine mixed with fluid from anal glands (Schaller 1972)
        • Many compounds in lion urine (Andersen and Vulpius 1999)
          • Communicative function unknown
      • Spray onto bushes, rocks, tree trunks, small hills, at water holes (Wilson and Mittermeier 2009; Estes 2012)
        • Also along roads and paths
    • Feces used much less frequently (Wilson and Mittermeier 2009)
  • “Scuffing” or scraping (Estes 2012)
    • Performed by both sexes
    • Scrape/rake ground with hind feet
      • Repeated from 2-30 times or more (Schaller 1972)
      • May or may not do while urinating
    • Glands between toes leave scent (Wilson and Mittermeier 2009)
    • Will also use claws to rake the ground and trees (Schaller 1972; Estes 2012)

Walk and run

• Walking speed (Schaller 1972; Rudnai 1973)
  • 3-4 km/hr (1.86-2.49 mph)
• Running
  • Bursts of speed (Schaller 1972)
    • Lack stamina to pursue prey over long distances
  • Maximum speed
    • Can run up to 45-60 km/h (28-37 mph) for short bursts (Schaller 1972; Chassin et al. 1976; Wilson and Mittermeier 2009)
      • Usually not more than 100-200 m

See Diet and Feeding

Food competitors (Schaller 1972, except as noted)

• Compete with scavengers for kill remains
  • Hyenas, jackals, vultures
    • May withdraw at lions’ approach and wait nearby for leftovers
    • Lions and hyenas exhibit reluctance to associate with each other
    • Jackals and vultures demonstrate more caution around lions than hyenas do
• Lions steal food from other predators
  • Take kills made by hyenas, leopards, and cheetahs
  • Leopards take kills into trees to avoid confrontations with lions (Wilson and Mittermeier 2009)

Grooming and social licking strengthen the social bonds between pride members.

Here, a lion cub is licked by her mother.

Image credit: © San Diego Zoo Wildlife Alliance. All rights reserved.

This play behavior, called "wrestling," is commonly observed in cubs 2 to 6 months old.

Cubs bound, chase, and wrestle for extended periods. They also play with objects, such as sticks and tufts of grass.

Image credit: © San Diego Zoo Wildlife Alliance. All rights reserved.