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

Activity Cycle

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

Seasonal movements and dispersal

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)

Territory

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

Social Groups

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)

Aggression

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 

Play Behavior

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)

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)
  • 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)

Communication

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)

Locomotion

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

Hunting Behaviors

Interspecies Interactions

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)

Morning Bath

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.

Wrestling Match

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.

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