African Elephants (Loxodonta africana and L. cyclotis) Fact Sheet: Behavior & Ecology

Daily patterns

• Savanna elephants
• Active day and night, but usually rest during hottest times of day (Guy 1976; Leggett 2010; Nowak 2018)
• More active at night during dry seasons and in dry areas (Loarie, Aarde, et al. 2009), and where poachers or human disturbance occur (Von Gerhardt et al. 2014; Gaynor et al. 2018; Ihwagi et al. 2018)
• Forest elephants
• Early research suggests activity during the day and at night (Wrege et al. 2012; Turkalo et al. 2013; Kely et al. 2019; Reyna-Hurtado et al. 2023)
• Some studies suggest feeding until about 14:00 and greater variety of behavior—excavating, bathing, fighting, sleeping—in the afternoon (Nowak 2018)
• In one study, moved between forest and grasslands at dawn and dusk (Mills et al. 2018)
• Avoid poachers and human activity during the day, similar to savanna elephant (Wrege et al. 2010; Beirne et al. 2021)

Feeding

• Accounts for majority of wakeful activity (Beekman and Prins 1989)
• Foraging in the morning thought to help elephants conserve energy and reduce heat stress (eg, Du Plessis et al. 2021)
• Savanna elephant
• Generally, at least half of active time spent feeding (eg, Wyatt and Eltringham 1974; Lindsay 2011; see literature reviewed by Sach et al. 2019)
• May be up to 80%, when less food available (e.g., dry season, drought years)
• Forest elephant
• Thought to forage mainly during the day (Blake 2002; Turkalo and Barnes 2013)
• Also see Diet & Feeding

Resting and sleeping

• Savanna elephant
• Generally, rest for short periods several times a day (eg, Wittemyer et al. 2017)
• Sleep for relatively brief periods (typically about 1 to 2 hours), for a total of about 3 to 5 hours in each 24-hour period (Eltringham 1982; Estes 1993; Wittemyer et al. 2017)
• Commonly rest/sleep late at night, between midnight and dawn, and during the early afternoon (hottest time of day) (eg, Wittemyer et al. 2017; Nowak 2018)
• Desert populations may have distinct nighttime rest periods (Leggett 2010)
• May sleep at night lying down, but usually rest standing up during the day (Eltringham 1982; Poole et al. 2013)
• For groups, sleeping and waking often synchronized within the herd (Eltringham 1982)

Human impacts

• Today, elephant ranges and movements mainly structured by human settlements and land use (including managed protected areas) (eg, Blake et al. 2008; Molina‐Vacas et al. 2020; Gobush et al. 2021; Leimgruber and Songer 2021; Wall et al. 2021)
• Green forage, surface water, and other environmental factors influence elephant space use within range constraints exerted by humans (Young et al. 2009; Wall et al. 2021; Benitez et al. 2022)

Home range

• Savanna elephants
• Annual range size of roughly 100 to 300 km², according to a continent-wide study by Wall et al. (2021)
• Largest home ranges observed in desert-dwelling elephants (eg, Wall et al. 2013)
• Males and females typically use distinct areas, except during breeding (see Poole et al. 2013)
• Move to find water and high-quality forage (Bohrer et al. 2014; Nowak 2018)
• Elephants constantly learn and recall detailed knowledge of landscapes (Polansky et al. 2015 and see Presotto et al. 2019)
• Forest elephants
• Annually, range over an area of less than 100 km², up to 2,000 km² (Turkalo and Barnes 2013; Beirne et al. 2021)
• May move more during the wet season, while seeking out preferred fruits (Mills et al. 2018; Beirne et al. 2020)
• Home ranges of females slightly smaller than males (Beirne et al. 2021; Wall et al. 2021)
• More study needed, but at least some individuals appear to continually shift and explore their home range (Turkalo et al. 2013; Beirne et al. 2021)
• Similar to savannah elephant (eg, Goldenberg et al. 2018)

Migration

• Savanna elephant
• Various movement patterns reported in different regions, depending on amount of food and water available (Gobush et al. 2022)
• Resident, migratory, semi-migratory, or nomadic-like movements
• Some herds show strong fidelity to their past wet or dry season range areas (eg, Burton-Roberts et al. 2022)
• May move long distances each year in search of green vegetation and water (Bohrer et al. 2014; Tshipa et al. 2017; Nowak 2018)
• Do not move far if food, water, and shade remain available across seasons (Nowak 2018)
• Artificial (human-made) water holes expand elephants’ dry season range (see Loarie, Aarde, et al. 2009) and allow some elephants to remain resident year-round (Tshipa et al. 2017)
• During drought, experienced matriarchs use their knowledge of water and food sources to lead herds across long distances (eg, Foley et al. 2008; Byrne and Bates 2011; also see McComb et al. 2001)
• Similar leadership roles recently discovered among male elephants (see O’Connell-Rodwell et al. 2024)
• May use environmental cues, such as rainfall, to time movements as the wet season begins (Kioko et al. 2020; Pandraud et al. 2022)
• Forest elephants
• Seasonal movements influenced by temperature and rainfall, despite living in wet forests (Beirne et al. 2020; Beirne et al. 2021)
• Tend to use similar areas from year to year, though males may be more migratory and females more resident (Turkalo and Barnes 2013; Turkalo and Fay 1995; Turkalo et al. 2013; Molina‐Vacas et al. 2020; Beirne et al. 2021)

Dispersal

• Savanna elephants
• Males (Vidya and Sukumar 2005; Poole et al. 2013 citing Lee and Moss 1999)
• Near puberty, leave birth family for extended periods to socialize with other families (Evans and Harris 2008; Lee, Poole, et al. 2011; Nowak 2018)
• Later, disperse and leave birth family permanently
• Full independence reached gradually (Lee and Moss 1999)
• Process can take 5 months to as long as 8 years [Amboseli population] (Lee, Poole, et al. 2011)
• On average, occurs at around 13 to 14 years old—but ages range from 7 to 19 years old [Amboseli, Samburu populations] (Lee, Poole, et al. 2011; Wittemyer et al. 2013)
• Females
• Continue living with birth family, though might disperse as a family group, singly, or in female pairs—eg, to recolonize unoccupied habitat areas (Ahlering et al. 2012)
• Forest elephants
• Both sexes gradually separate from birth family (Turkalo et al. 2018)
• Female forest elephant behavior very different from female savannah elephant, where daughters usually remain with herds that include their mothers, sisters, and grandmothers (matriarchal groups) (Turkalo et al. 2018)
• Possible social strategy to prevent competition between offspring and their parents (Turkalo et al. 2018; Goldenberg et al. 2021)
• Daughters may disperse when ready to give birth to their first calf or when mother dedicates increased care to younger siblings (Turkalo et al. 2018; Goldenberg et al. 2021)
• Age at independence
• About 15 years old (median age for both males and females) (Turkalo et al. 2018)
• But some individuals may separate from mother or conceive at younger ages (Turkalo 2013; Turkalo et al. 2017; Turkalo et al. 2018)
• Reach full independence months or years after initial separation (Turkalo et al. 2017; Turkalo et al. 2018)

Home ranges commonly overlap

• Not territorial (both species) (Shoshani 1992; Poole et al. 2013; Nowak 2018)
• Generally, home ranges overlap, especially those of family groups (Douglas-Hamilton 1972; Nowak 2018)
• But groups may segregate by social rank or use home ranges with less overlap to avoid competing for resources (eg, during drought) (eg, Wittemyer et al. 2007; Blake and Maisels 2023)

Social structure

• Fission–fusion social structure in both species (eg, Moss and Poole 1983; Wittemyer et al. 2005; Archie et al. 2006; Wittemyer et al. 2009; Moss and Lee 2011b; Fishlock and Lee 2013; Head et al. 2013; Poole et al. 2013; Athira and Vidya 2021)
  • Members of a group change often, coming together to form larger groups and splitting apart
    • Fusing more common when food more abundant
    • Splitting more common when food is scarce
  • Dynamics influenced by many factors, including social bonds, season, home ranges, habitat, reproductive state, and genetic relatedness
  • Excellent memory for recognizing family members and distinguishing between familiar/unfamiliar individuals
• Savanna elephants
  • Adults usually split into same-sex groups (Moss and Poole 1983; Kioko et al. 2020)
    • Adult males usually associate with females for only short periods
    • Also see “Social associations among males,” below
  • Herd behavior is highly gregarious (eg, Douglas-Hamilton 1972; Lee 1987; McComb et al. 2011; Nowak 2018)
    • Group members cooperate to find resources, defend the herd, and care for calves (females)
      • Also see Defense and Parental Care
  • Females form complex social bonds—among the most complex of any animal (eg, Moss and Poole 1983; Moss and Lee 2011b; Goldenberg et al. 2016)
    • Solitary females very rare (Nowak 2018)
  • Groups tend to consist of adult females—primarily female relatives and their offspring (e.g., Douglas-Hamilton 1972; Douglas-Hamilton 1972 citing Laws and Parker 1968; Archie et al. 2006; Moss and Lee 2011b; Charif et al. 2015)
    • Led by matriarch
    • Often multi-generational, where grandmothers, and their daughters and granddaughters live together
  • Usually about 10 to 25 individuals in a core group (family group) (Kingdon 1979; Young et al. 2009; McKnight 2015)
    • But have relationships with extended family, so groups may become several times larger—more than 50 individuals (Owens and Owens 2009; Moss and Lee 2011b; Poole et al. 2013)
    • Mixed groups (eg, breeding groups) may include 1 or more adult males (eg, McKnight 2015)
    • Where poaching occurs, a “group” may be reduced to a single female and her one offspring (Owens and Owens 2009; Moss and Lee 2011b)
  • Affiliations beyond extended family groups known as “clans” (Laws 1969; Moss and Poole 1983; Wittemyer et al. 2005; Archie et al. 2011; Poole et al. 2013; Charif et al. 2015; Nowak 2018)
    • For detailed information about core, bond, and clan groups, see Wittemyer et al. (2005) and review by Athira and Vidya (2021)
  • Large, temporary aggregations in open habitats can reach into the hundreds of individuals (Moss 1988; Owens and Owens 2009; Poole et al. 2013; McKnight 2015; Nowak 2018)
• Forest elephants
  • Very different social organization than savanna elephant—more similar to Asian elephants, which also commonly live in forests (eg, White et al. 1993; Turkalo and Fay 1995; Turkalo et al. 2017; Goldenberg et al. 2021; Blake and Maisels 2023)
    • Difference may be due to patchy food or difficulty of traveling through dense forest in larger groups
    • Social role of matriarchs limited or absent outside of small family groups, which are comprised of mothers and their offspring
      • No extended matriarchal hierarchies, as in savanna elephants
  • Females typically observed in small groups of 2 to 5 individuals, but also commonly observed singly (White et al. 1993; Turkalo and Fay 1995; Turkalo et al. 2013; Nowak 2018; Reyna-Hurtado et al. 2023)
    • Groups commonly comprised of 1 or 2 related females and their dependent young (White et al. 1993; Turkalo and Fay 1995; Turkalo et al. 2013; Turkalo and Barnes 2013; Goldenberg et al. 2021; Reyna-Hurtado et al. 2023)
  • Core groups have stable associations over time (Goldenberg et al. 2021; Blake and Maisels 2023)
    • Uncommonly, 2 or 3 core groups may temporarily fuse (Head et al. 2013)
  • Sometimes aggregate in natural forest clearings and at mineral springs (Turkalo and Fay 1995; Fishlock and Lee 2013; Turkalo et al. 2013; Turkalo and Barnes 2013; Goldenberg et al. 2021)
    • Larger family units of up to 20 individuals observed at Dzanga clearing, Central African Republic (Turkalo and Barnes 2013)
  • Adult males mainly associate with females during breeding (Turkalo et al. 2013; Nowak 2018)

Social associations among males

• Savanna elephants
  • Solitary males common (Chiyo et al. 2011; Nowak 2018)
  • Some males not very sociable (Poole 1987; Ahlering et al. 2012; Nowak 2018), some associate with unrelated males, and others still more highly bonded (O’Connell-Rodwell et al. 2024)
    • Males (not in musth) can have temporary, yet sometimes strong social ties (Goldenberg et al. 2014; Nowak 2018; O’Connell-Rodwell et al. 2024)
    • Other adult males maintain stable social relationships (in connected networks) over long time periods (Murphy et al. 2019; O’Connell-Rodwell et al. 2024)
  • Group sizes
    • McKnight (2015) reported groups of up to 18 males
  • Older males—or those more integrated or dominant in a group’s social network—strengthen associations among group members (Chiyo et al. 2011) and are more likely to lead group movements (Allen et al. 2020; O’Connell-Rodwell et al. 2024)
    • Younger males—those without in-depth spatial and ecological knowledge—thought to follow and gain ecological knowledge from experienced, adult males (Allen et al. 2020)
• Forest elephants
  • Juvenile and subadult males associate temporarily for a day to several days (Turkalo and Barnes 2013)
  • Adult males usually live singly, outside of breeding periods (White et al. 1993; Turkalo and Fay 1995; Fishlock and Lee 2013; Head et al. 2013; Head et al. 2013; Turkalo and Barnes 2013; Nowak 2018)
    • Rarely form bachelor groups (of only 2 to 3 individuals)

Dominance hierarchies

• Savanna elephants
  • Strong linear dominance hierarchies (Archie et al. 2006; Wittemyer and Getz 2007; de Silva et al. 2017)
  • Individual and group dominance structured by age (older = typically more dominant) (eg, Athira and Vidya 2021)
    • Older, larger females dominant to smaller, younger females (Archie et al. 2006; Nowak 2018)
    • Larger families with older matriarchs dominant to smaller family groups (Mutinda et al. 2011)
      • Have better access to preferred habitats and water, particularly during the dry season (Wittemyer et al. 2007)
  • Males in musth, especially those of larger body size, dominant to males not in musth (eg, Poole and Moss 1981; Hall-Martin 1987; Poole 1987; Poole 1989a)
    • Presence of adult males regulates behavior of younger males (eg, less defensive/aggressive and less fearful) (eg, Slotow et al. 2000; Allen et al. 2021)
  • Males form linear hierarchies when resources (eg, water) are limited (O’Connell-Rodwell et al. 2011)
• Forest elephants
  • Males confront other males in forest clearings and at waterholes to establish/maintain their dominance status (Turkalo and Fay 1995; Turkalo and Barnes 2013; Nowak 2018)
  • Adult males more dominant than females and juvenile males (eg, at waterholes) (Turkalo and Fay 1995)
    • When present, tend to control access to preferred waterholes and minerals for long periods (Turkalo and Fay 1995; Wrege et al. 2012)

Orphaned calves—challenges

• Calves unlikely to survive if orphaned before reaching 2 years old (Poole and Cobb 1989; Poole and Thomsen 1989; Moss and Lee 2011; Parker et al. 2021)
• Survivors may or may not become fully integrated into a new social group (Goldenberg and Wittemyer 2018)
• Weaker social bonds with adult females can result in orphans having less access to resources, such as food, water, or shade (Goldenberg and Wittemyer 2017; Goldenberg and Wittemyer 2018)
• May not gain social or nuanced survival knowledge typically learned from members of their birth family (Goldenberg and Wittemyer 2017; McComb, Shannon, et al. 2011; Shannon et al. 2022)
• Orphans may be subjected to more aggression from group members, compared to non-orphans, which likely causes more stress (Goldenberg and Wittemyer 2018)
  • However, some elephants may be able to adapt to prolonged stress (see Parker et al. 2022; Pokharel and Brown 2023)
• Stunted growth observed among orphans of one Kenyan population (Parker and Wittemyer 2022)

Play behavior

• Observed from infancy through adulthood, though declines over a lifetime (Lee and Moss 2014)
• Elephants curious and quick to investigate (Poole and Granli 2011)
• May use all parts of body—trunk, mouth, tusk, feet, etc.
• Helps juveniles develop physical abilities, as well as social skills (eg, Evans and Harris 2008; Lee and Moss 2014)
• Females: develop protector and leadership roles within families; play with younger calves
• Males: engage with non-relatives; develop social relationships
• Both gentle and “rough-and-tumble” play observed (eg, Evans and Harris 2008; Poole and Granli 2011; Lee and Moss 2014)
• Wrestle, shove, and spar
• Splash and spray water
• Roll in mud
• Mock charge and chase other elephants
• Manipulate objects or lift/thrash vegetation
• Give trumpet sounds (Poole 2011)
• Young elephants chase mammals and birds (Poole et al. 2013)

Grief and consolation behaviors

• Elephants investigate, approach, smell, and touch other dead elephants, including relatives and non-relatives (eg, Moss 1992; Payne 2003; Douglas-Hamilton et al. 2006; Poole and Granli 2011; Hawley et al. 2018; Goldenberg and Wittemyer 2020; Stephan et al. 2020)
• Some other example responses include social behaviors, nudging or lifting behaviors, guarding the body, fear-related behaviors, picking up bones (Moss 1992; Poole 1996; Payne 2003; Douglas-Hamilton et al. 2006; Goldenberg and Wittemyer 2020; Stephan et al. 2020), attempting to feed an elephant that died, or covering the dead body with dirt or vegetation (Moss 1992; Poole and Granli 2011)
• Behavior varies greatly from individual to individual
• Lifting the body and vocalizations reported only when an elephant had recently died (Goldenberg and Wittemyer 2020)
• Elephants may interact with dead individuals for extended periods, particularly using their highly developed sense of smell (Goldenberg and Wittemyer 2020)
• Show interest in other kinds of dead animals (Poole et al. 2013)
• Some individuals engage more with elephant remains vs. non-elephant remains (McComb et al. 2006; Goldenberg and Wittemyer 2020)

Tactile

• Family members communicate through touch
  • May rub with a foot or slap with the trunk, often while standing
  • Uses of the trunk
    • Trunk used in greeting
      • May be held out to an approaching elephant as a greeting
      • A lower-ranking animal will insert its trunk tip into the other’s mouth
    • Other uses of the trunk
      • An individual will caress another
      • Mother may guide her calf by gripping its tail with her trunk
      • Two individuals may intertwine trunks
      • Trunk used to check reproductive status

Vocalization

• Click here for audio of the African Elephant. Provided by The Cornell Lab of Ornithology Macaulay Library.

• Sound diversity
  • 4 different sounds are produced
    • Each is distinguished by gradations in pitch, duration, and volume
  • Rumbling
    • Frequencies variable; most are below human hearing
    • Functions
      • Important to sexual behavior; imparts distance information
        • Louder sounds may carry over several kilometers
      • A signal to more, or a warning
      • Softer rumbles may promote group cohesion
        • Quiet, though audible calls, heard during feeding events
      • Social greeting
        • Rumble or growl may be given as a greeting
      • Elevated, more bellowing rumbles given when in fear or pain
  • Trumpeting
    • Blowing through nostrils
    • Indicates excitement
  • Squealing
    • Juvenile distress call
  • Screaming
    • Adult distress call
    • Often in conjunction with trumpeting
    • Possibly meant to to intimidate another
• Infrasonic communication
  • Discovered by Katherine Payne, based on observations of elephants raised under managed care made in 1984
• Species specific characteristics
  • Savannah/bush elephants vocalize at 14-24 hertz
  • Forest elephants vocalize around 5 hertz

Olfaction/Scent Marking

• Scent glands
  • Temporal gland secretions "musky"
    • Secretions is under autonomic control
    • Produced when an individual is excited or anxious

Walking

• Adults generally walk (McKay 1973)
  • Only run when fleeing danger or charging
  • Move legs on same side of body, even at higher speeds (Hutchinson et al. 2003; Hutchinson et al. 2006)
    • Left hind, left front; right hind, right front (Hutchinson et al. 2006)

• Basic gait
  • 3 feet on the ground most of the time
  • At least 1 foot always on ground
  • Body has a rolling motion and head bobs during fast walk

• Speed
  • Slow walk
    • About 0.5 kph (0.3 mph) or slower (Ngene et al. 2010; Chamaillé‐Jammes et al. 2014; Troup et al. 2020)
      • When eating, resting, socializing in one location (Ngene et al. 2010)
    • 2 to 3 kph (1 to 2 mph) (Hutchinson 2006)
      • When shifting between resource patches
  • Faster walk
    • 6 to 8 kph (4 to 5 mph) (Estes 2012)

“Running”

• Unusual hybrid gait for moving fast (Hutchinson et al. 2003 in Asian elephant)
  • Legs support bouncing motion, and joints bend more as speed increases (Hutchinson et al. 2003; Hutchinson et al. 2006)
• Primarily increase speed by taking more strides (Hutchinson et al. 2006; Estes 2012)
• Maximum speed
  • Up to about 26 kph (16 mph) (Hutchinson et al. 2003; Ngene et al. 2010; Estes 2012)

Jumping

• Cannot jump (Estes 2012)
• Have difficulty crossing some ditches/trenches
  • Generally unable to cross gaps wider than about 2 m (7 ft)

Other postures

• Able to kneel, squat, roll, and sit on haunches (eg, when getting up and lying down) (Estes 2012; Poole et al. 2013)
• Can balance back legs to reach browse up high (Estes 2012)

Herbivore competitors

• Coexist with 3 other large mammal species (Eltringham,1982)
• Buffalo (Syncerus), bush pig (Potamochoerus), bushbuck (Tragelaphus scriptus)
• Elephants typically ignore other herbivores
• Such as antelopes, warthogs and zebras

Carnivore predators

• Elephants seek out lions in their vicinity to chase them away

Elephants as "keystone" species

• Elephants impact on animals and plants
• Elephants may stop the progression of grassland to forest or shrub lands
• Grasses support grazing ungulates which are in turn food for carnivores
• Elephants maintain trails used by all species
• Elephants keep down vegetation around water holes
• Process creates safe drinking spots for all species

Elephant calf playing in the mud.

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

Caloi (1992)
Estes (1991)
Estes (1993)
Eltringham (1982)
Moss (2000)