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African Elephants (Loxodonta africana and L. cyclotis) Fact Sheet: Reproduction & Development

Courtship

Mating vocalizations

  • Females
    • Females in estrus emit low-frequency calls (“estrus rumbles”) to attract males over long distances (Poole et al. 1988; Langbauer et al. 1991; K.M Leong et al. 2003; Garstang 2004)
      • Signals their reproductive receptivity to males
      • Call structure helps males judge direction and distance to travel
    • Respond to male “musth rumbles” with loud, low-frequency vocalizations (Poole 1987)
  • Males
    • Low-frequency calls (“musth rumbles”) given by breeding males while searching for females in estrus (Poole 1987; Poole et al. 1988; Langbauer 2000; Stoeger and de Silva 2014)
      • Advertises a male’s reproductive and dominance status (Poole 1999; LaDue et al. 2022)
      • May also help musth males avoid one another (Poole et al. 1988)

Musth (in males)

  • Unique sexual state in breeding male elephants (Jainudeen et al. 1972; Poole and Moss 1981; Poole 1987; LaDue et al. 2022, and as noted)
    • Period of heightened sexual activity and aggressive behavior in breeding males (Hall-Martin 1987; Poole 1987; Poole 1989a)
  • Functions and adaptive value (Hall-Martin 1987)
    • Allows males, especially older males, entry into matriarchal groups
      • Signals to females their readiness to breed
    • Facilitates female mate choice
    • Reduces dominance-related aggression among adult males, often eliminating the need for physical combat
      • Helps prevent injury and death
  • Physical indicators (Poole and Moss 1981; Hall-Martin and Van Der Walt 1984; Hall-Martin 1987; Poole 1987)
    • Glands on sides of head (temporal glands) swell and secrete a dark, oily liquid
    • Dribble urine between hind legs
    • Additional sex pheromones present in breath
    • Pungent body odor
  • Behavioral indicators
    • Increased movement or “roaming” (Moss 1983; Hall-Martin and Van Der Walt 1984; Hall-Martin 1987; Taylor et al. 2020)
    • Sexually active males smell and taste urine of females to assess reproductive condition (Moss 1983; Hall-Martin 1987)
    • Male associates more with females and guards breeding females (Poole and Moss 1981; Moss 1983; Hall-Martin 1987; Poole 1987, Poole 1989b)
    • Increased aggressive behavior (Poole and Moss 1981; Hall-Martin 1987; Poole 1987; Poole 1989a)
      • Musth males will confront, threaten, chase, shove, and fight other males (Moss 1983; Poole 1989a)
      • Smaller males remain on periphery of female-led group and avoid encounters with larger males
    • More frequent displays (e.g., musth walk, ear waving/flapping) (Poole 1987; LaDue, Vandercone, et al. 2022b) and breeding-specific vocalizations (low, pulsing “musth rumble”) (Poole 1987)
  • Savanna elephants
    • Timing
      • Musth can occur at any time of year in a population (eg, Poole and Moss 1981)
      • Not all breeding males enter musth at the same time, but individual males begin musth at about the same time every year (Poole and Moss 1981; Poole 1987; Poole 1989a)
      • In some populations, timing appears to be influenced by a combination of season and age/social status (Chase LaDue, personal communication, 2025)
        • Ex. Older, more dominant males tend to be in musth during rainy season (when resources are abundant) and younger, less dominant males enter musth during the dry seasons
    • Age of first musth
      • Usually by about 25 to 30 years old (range 17 to 34 years) (Poole 1987; Poole et al. 2011; Poole et al. 2013: 201)
    • Duration increases with age (Poole 1987; Poole et al. 2011)
      • Typically lasts 4 to 8 weeks in males 36 to 40 years old; up to 10 weeks in males 46 to 50 years old (Poole et al. 2011)
      • Smaller, low-ranking males may come into musth but only for short, sporadic periods (Poole 1987)
    • Thought to spend more time foraging and restoring body weight during non-musth periods (Taylor et al. 2020)
      • Most males lose weight during musth, due to increased activity and likely an increased metabolism (from higher levels of androgen hormones, like testosterone) (Poole 1989a)
  • Forest elephants
    • Further study needed (especially apart from Dzanga population) (Chase LaDue, personal communication, 2025)
    • Forests have less seasonal change in resources (such as food), so patterns may be more similar to Asian elephants—which also live in forests—than savanna elephants (Chase LaDue, personal communication, 2025)
    • At Dzanga Bai, available data suggest that musth is synchronized (Turkalo and Barnes 2013; Nowak 2018 citing works by A. Turkalo and colleagues)
      • Males enter musth at about the same time each year, most commonly during drier months
    • Musth observed in only the largest males at Dzanga Bai (Turkalo and Barnes 2013)
    • Forest clearings may facilitate male–male competition (Fishlock and Lee 2013; Blake and Maisels 2023) or other aspects of mate choice

Mating system and mate choice

  • Female mate preference
    • Tend to mate with large males in musth (Moss 1983; Poole 1989b)
      • Males 45 to 50 years old most successful at siring young (especially compared to younger males)
    • Older, experienced females appear to exert more mate choice (Moss 1983)
  • Males compete for access to females (eg, Moss 1983)
    • Females may attract several males with their calls and scents, but may only mate with a dominant bull during their short conception period (eg, Moss 1983; Hildebrandt et al. 2011)
    • Older males in musth likely better able to outcompete rival males (Poole 1989b)
    • While guarding females, musth males chase away younger males and prevent them from mating (Poole 1989b; Poole et al. 2013)

Courtship behavior and copulation

  • Before mating
    • Female appears wary and watchful (Moss 1983; Poole et al. 2013)
      • May avoid males for hours or days before soliciting attention of certain males (Moss 1983)
    • Female then initiates an “estrus walk” (Moss 1983)
      • Female moves away from herd with one male (or less commonly, 2 or 3 males)
      • Near musth male, holds head high and walks in circles or figure-8s
    • Chase behavior often occurs in later stages (Moss 1983)
  • Copulation (Moss 1983, except as noted)
    • Female stands still and braces with forelegs to enable male to mount
    • Mating may occur several times per 24-hour period in savanna elephants (Poole et al. 2013), or a couple times over about 2 days in forest elephants (Turkalo and Barnes 2013)
  • After mating
    • Female stands still or sometimes runs off
    • In savanna elephant, female that just mated may flap ears and give rumbles, which are joined by rumbles, trumpets, and roars of nearby elephants—usually family members (Moss 1983; Poole et al. 1988; Poole 1989b; Poole 2011; Poole et al. 2011)
    • Male may guard females in a herd (eg, Moss 1983; Archie et al. 2007; Turkalo and Barnes 2013)

Reproduction

(Note: for wild populations, unless noted)

Sexual maturity

  • Savanna elephant
    • Females
      • For Amboseli and Addo populations, mature by about 12 years and give birth to first calf at about 14 years (Moss 2001; Gough and Kerley 2006; Freeman et al. 2009; Moss and Lee 2011; Lee et al. 2016)
        • Other studies found similar ages at first birth of 14 to 16 years old, on average (Owens and Owens 2009; Poole and Granli 2022)
        • Females can give birth at younger ages in populations that experience heavy poaching (Owens and Owens 2009; Nowak 2018)
    • Males
      • Begin producing sperm at about 10 to 14 years old (Nowak 2018)
      • Majority of males experience their first musth by age 30 (Poole 1987; Poole et al. 2013)
      • In the wild, not of sufficient social status to compete and mate until into their 20s (Poole 1989a)
        • Access to breeding females limited by more dominant males
        • Most males under 25 lack experience and female cooperation to mate successfully (Poole and Thomsen 1989)
    • In managed care, sexual maturity and breeding behaviors can begin at much younger ages, perhaps due to good nutrition (reviewed by Brown 2014, 2019)
      • Affects growth and reproductive development, leading to earlier maturity (as young as 5 to 10 years in U.S. zoos)
  • Forest elephant
    • Females
      • Mature at about 12 to 14 years old (Turkalo and Barnes 2013)
        • But much variation reported
        • Some studies find some individuals giving birth as young as 8 to 10 years old (Theuerkauf et al. 2000; Turkalo et al. 2017)—yet, on average, females in Dzanga (Central African Republic) do not begin giving birth until their early 20s (Turkalo et al. 2017; Turkalo et al. 2018)
        • Reproductive activity may be limited by ecological resources (Turkalo et al. 2017)
    • Males
      • Likely physically mature at about same age as females—but like savanna elephants, begin successfully mating at an older age (Turkalo and Barnes 2013)

Breeding

  • Seasonality
    • Savanna elephant
      • Can occur during any time of year, and may coincide with seasonal or annual cycles of high rainfall (Smuts 1975; Hall-Martin 1987; Sukumar 2003; Brown 2014; Wittemyer, Barner, et al. 2007; Poole et al. 2013; Nowak 2018)
        • Influenced by availability of high-quality forage (eg, Wittemyer, Barner, et al. 2007)
        • Several conception peaks possible (eg, Freeman et al. 2009) in places where vegetation is available throughout much of the year (Hall-Martin 1987)
      • Females generally breed when in good body condition and when ample food available (Moss 2001; Wittemyer, Barner, et al. 2007; Lee, Lindsay, et al. 2011; Poole et al. 2013)
    • Forest elephant
      • Estrus peaks reported during 2 dry seasons (Turkalo and Barnes 2013), but females can enter estrus and conceive during any time of year (Turkalo et al. 2018)
      • Most calves born from May to August, in the middle of the wet season (Turkalo et al. 2018)
  • Reproductive cycles
    • Females
      • Polyestrous (Nowak 1999; Hildebrandt et al. 2011; Brown 2019)
      • Long estrus cycle but short conception period (Moss 1983; Rasmussen and Schulte 1998)
      • Estrus cycle
        • About 15 to 17 weeks, on average (Plotka et al. 1988; Brown 2000; Wittemyer 2011a)
          • Range: 13 to 17 weeks, according to Brown (2014, 2019)
      • Peak estrus period (conception period)
        • Typically, 4 to 6 days in savannah elephant (Moss 1983; Poole et al. 2013)
        • Thought to be similar in forest elephants (“several days” long, according to Turkalo and Barnes 2013)
    • Males
      • Can mate at any time of year (Jainudeen et al. 1972; Hall-Martin 1987)
        • Do not need to be in musth, though large males in musth dominate mating opportunities
  • Lifetime reproduction
    • Females (both species)
      • Fecundity generally consistent until age 40 or 45, then declines slightly (Moss 2001; Freeman et al. 2013; Poole et al. 2013; Lee et al. 2016; Turkalo et al. 2018)
        • Some females 50–60 years or older can still calve (Moss 2001; Poole et al. 2013; Lee et al. 2016; Turkalo et al. 2018), but most “grand matriarchs” have finished reproducing (Freeman et al. 2009, 2011, 2013)
          • Social and ecological knowledge important to group survival (eg, McComb et al. 2011)
    • Males
      • Males also capable of reproducing at age 60 (Poole et al. 2011)

Gestation and Birth

Gestation

  • Savanna elephant
    • About 22 months, on average (Perry 1953; Poole 1989a; Meyer et al. 2004; Nowak 2018; reviewed by Brown 2014, 2019)
      • Range: 17 to 25 months
  • Forest elephant
    • About 22 months, on average (Shoshani and Tassy 2013; Turkalo and Barnes 2013)

Birth

  • Number of young
    • Usually one calf (both species) (Shoshani and Tassy 2013; Turkalo and Barnes 2013)
      • Twins very rare (Moss et al. 2019)
  • Weight at birth
    • Savanna elephant
      • About 90 to 120 kg (200 to 270 lb), on average (Nowak 1999; Nowak 2018)
  • Shoulder height at birth
    • Savanna elephant
      • About 80 to 90 cm (31 to 35 in) (Amboseli, Kruger populations) (Lee and Moss 2011, Table 14.2; Poole et al. 2013)
    • Forest elephant
      • About 60 cm (24 in) (Turkalo 2013, Table 1)
  • Interbirth interval
    • Savanna elephant
      • Typically, about 3 to 5 years [Amboseli, Addo, and Kruger National Park populations] (Moss 1983; Gough and Kerley 2006; Poole et al. 2013; Freeman et al. 2009; Lee et al. 2016)—but varies between and within populations (Sukumar 2003; Freeman et al. 2009; Poole et al. 2013)
        • Range: about 3 to 9.5 years (Eltringham 1982; Sukumar 2003; Poole et al. 2013 citing Viljoen 1988)
    • Forest elephant
      • About 5 years, according to Turkalo et al. (2017, 2018) [Dzanga Bai (Central African Republic)]
        • Range: about 1.5 to 11 years (Turkalo et al. 2018)
    • Food availability, population density, and death of a calf can all influence how often a female gives conceives and gives birth
  • Labor (savanna elephant; Poole et al. 2013, except as noted)
    • Females often give birth at night
    • Family members usually nearby
    • Usually stand but sometimes lie down during labor
    • Mother and other females gently remove birth sac
    • Newborn able to stand after about 15 minutes
    • Family members vocalize, show interest in newborn calf, carry and toss placenta, and follow the mother and newborn
    • Newborns stay close to mother/family members (Turkalo and Barnes 2013) and begin moving with herd after about 2 days (Nowak 2018)

Parental Care

Investment in care

  • Allomaternal care (“allomothering”) (Lee 1987; Lee and Moss 2011; Turkalo and Barnes 2013)
    • Calves looked after by their mothers and other adult and subadult females in their herd (Gadgil and Nair 1984; Santiapillai et al. 1984; Rapaport and Haight 1987; Turkalo and Barnes 2013; Berliani et al. 2019)
      • Cooperative care yields a strong survival advantage for calves (Poole et al. 2013)
    • Females protect and reassure calves in a group
      • Guide or restrain calf movements using trunk, nudging with a foot, and/or gentle use of tusks (eg, helping a calf up a steep incline) (Lynn Von Hagen, personal communication, 2025)
    • Helpers also allow mother time to feed and rest
      • Important for milk production
      • Lactation puts high energy/food/nutritional demands on mothers
  • Role of grandmothers
    • Retain and “impart” knowledge to daughters and grand-calves (McComb, Shannon, et al. 2011; Mutinda et al. 2011)
      • Promotes social bonding and increases calf survival (Moss and Lee 2011; Lee et al. 2016)

Nursing

  • Mothers typically suckle one calf (Poole and Granli 2022)
    • Nursing more than one calf incredibly rare, as mother usually cannot produce enough milk for more than one calf (Moss et al. 2019)
      • But, in rare circumstances, reported for mothers that conceive again quickly after giving birth (eg, where intense poaching occurs (Owens and Owens 2009)
  • Male calves have higher energy demands compared to female calves (Lee and Moss 1986)
    • Longer time to birth of next calf when nursing a son
    • Producing enough milk can be a challenge for mothers during drought
      • A common cause of infant mortality (Lynn Von Hagen, personal communication, 2025)

Weaning

  • Savanna elephant
    • Early nutrition
      • Calves rely on milk exclusively for first 3 months (Wittemyer 2011c)
        • Then begin feeding on plants—but still heavily dependent on milk
      • Cannot use trunk to suck up/drink water until about 8 to 9 months old [savanna elephant] (Lee and Moss 2011)
        • Observed to get down on knees to drink water, until they gain enough trunk control (Lynn Von Hagen, personal communication, 2025)
    • Age at weaning
      • Calves depend on mother’s milk for 2 years (Wittemyer 2011c; Poole et al. 2013; Nowak 2018)
        • By 2 years old, diet is similar to adults
      • But most calves will continue to nurse as long as mother will tolerate suckling (Poole et al. 2013)
        • In Amboseli, full weaning at about 4.5 years (55 months) (Lee and Moss 2011)
        • Calf may be forced to wean after a younger sibling is born (Poole et al. 2013 citing Lee and Moss 1986 and Whyte 2001)
  • Forest elephant
    • Typical age at weaning not yet scientifically demonstrated
      • Likely corresponds with birth of a mother’s next calf, similar to savanna elephant (Shifra Goldenberg, personal communication, 2025)
    • Some calves observed suckling until 5 years old (Turkalo and Barnes 2013)

Life Stages

Note: See Turkalo (2013) for a detailed aging rubric for forest elephants.

Infant (less than 1 year old)

  • Newborn
    • Able to stand about 15 minutes after birth (Poole et al. 2013)
    • Walk is unsteady and “stiff-legged” (Turkalo 2013)
  • First 6 months
    • Body begins to fill out (Turkalo 2013)
    • Gradually walk more steadily (Turkalo 2013)
    • Learn to use trunk (Turkalo 2013)
    • Begin to feed on plants (Wittemyer 2011c; Turkalo 2013)
    • Stay close to mother, but begin to explore/feed/play short distances away from her (Lee and Moss 2011; Turkalo 2013)
  • 6 months to 1 year
    • Rapidly develop motor skills
      • Trunk coordination improves (Turkalo 2013)
    • Easily feed on vegetation (in addition to suckling) (Turkalo 2013)
    • Can now use trunk to drink (suck up) water (Turkalo 2013)
  • 1 to 2 years
    • At 1 year old, have full trunk coordination (Turkalo 2013)
    • After 18 months, permanent tusks erupt and replace tushes (deciduous tusks; also see Tusks) (Fowler 1993; Turkalo 2013)

Juvenile

  • Physical development
    • Grow fastest in first 10 years of life (Moss and Lee 2011)
    • Noticeable sex differences in forest elephants by 6 to 7 years old (Turkalo 2013)
      • Bodies of males larger and bulkier than females
      • Tusks of males thicker and longer
  • Behavioral and social development
    • Social skills for females and males initially similar, but develop sex-specific behaviors (Poole et al. 2013 citing Lee 1986 and Lee and Moss 1999; Turkalo 2013)
      • Females interact with family members to a greater degree, forming tight relationships
        • Allomother younger calves
      • Males become more independent
        • Explore farther
        • Play with young elephants from other families
        • Spar with other juvenile males
    • Also see Dispersal

Subadults

  • Females
    • Body proportions match those of an adult (Turkalo 2013)
    • Approach full adult height by about 25 years old (Turkalo 2013)
    • Female savanna elephants grow throughout their lifetime (Nowak 2018)
      • Female forest elephants stop growing at reproductive maturity
  • Males
    • Reach full height by about 40 years old (Wittemyer 2011c; Turkalo 2013, Table 3)
      • In forest elephant, rapid growth from 15 to 20 years old (Turkalo 2013)
    • Gain weight throughout their lifetime (Nowak 2018)

 

Typical Life Expectancy

Wild populations

  • Life expectancy varies, depending on population, sex, amount of poaching pressure, etc.
  • Typical life expectancy across regions needs more study, but likely about 30 to 40 years (e.g., Moss 2001; Lee et al. 2012; Lee, Fishlock, et al. 2016; see Lee, Lindsay, et al. 2011 for Life Tables)

Managed care

Mortality

Survival rates

  • Also see Threats to Survival for discussion of poaching and human–elephant coexistence
  • Protected vs. unprotected habitat areas
    • Deaths from “natural causes” (eg, starvation, dehydration) can be comparable to deaths from ivory poaching, if elephants not able to roam (ie, range is limited to a small protected area) to find enough food and water (esp. during drought conditions) (Mukeka et al. 2022)
  • Male vs. female survival
    • Mortality for males generally higher across lifespan (Moss 2001; Gough and Kerley 2006; Poole et al. 2013; Wittemyer et al. 2021; Mukeka et al. 2022)
  • Calf survival
    • More savanna elephant calves die from drought and predators, compared to forest elephants (Wittemyer et al. 2013; Turkalo et al. 2018)
      • Drought lowers survival for adult females and their calves (eg, Lee and Moss 1986; Lee et al. 2013; Lee et al. 2016)
        • Affects milk production/quality (Lee and Moss 1986; Shrader et al. 2010; Lee et al. 2013) and increases travel distances to find food/water, which may result in calves becoming lost or separated from their family (eg, Loveridge et al. 2006)
      • Orphaned calves experience lower survival rates, which can harm population growth/recovery (eg, Goldenberg and Wittemyer 2018; Parker et al. 2021)
    • Turkalo et al. (2018) reported very high survival rates (97%) for 2-year-old calves at Dzanga Bai, a well-protected population of forest elephants (Turkalo et al. 2018)

Predation

  • Predators of calves and juveniles
    • Savanna elephant
      • Lions (e.g., Loveridge et al. 2006; Power and Shem Compion 2009)
      • Spotted hyenas (Poole et al. 2013)
      • Leopards (Nowak 2018 citing Blake 2004)
    • Forest elephant
      • Not thought to have non-human predators, but rarely, possibly leopards (Turkalo and Barnes 2013)
  • Predators of adults

Diseases (non-comprehensive)

  • Anthrax
    • Observed in wild savanna elephants but not forest elephants (Turkalo and Barnes 2013)
    • Can affect entire populations (Poole et al. 2013 citing Lindeque and Turnbull 1994)
      • A disease that might contribute (alongside other factors) to limiting excessive population growth (ie, too large for an ecosystem to support) (see Foley et al. 2024)
  • Encephalomyocarditis virus (abbreviated EMCV) (Grobler 1995; Poole et al. 2013)
    • A potentially fatal viral infection in savanna elephant (Poole et al. 2013)
      • More than 60 elephants died in one outbreak in Kruger National Park (1993 to 1994) (Grobler 1995)
  • “Floppy trunk syndrome” (Poole et al. 2013; Landolfi and Terrell 2018)
    • Observed in savanna elephants
    • Rare, progressive nervous system condition leading to total paralysis of the trunk
    • Affected individuals have difficulty drinking and eating, leading to poor condition and possibly death
    • Cause unknown; possibly a toxin or pollutant
  • Bacterial infections that cause blood poisoning (Azeem et al. 2020; Foggin et al. 2023)
    • Transmission pathway(s) not yet known, but can cause mass mortality events (Foggin et al. 2023)
      • In 2020, 35 elephants died from a Pasteurella-like infection in Zimbabwe (Foggin et al. 2023)
    • An elephant’s immune system can be compromised by multiple stressors: drought, heat, malnutrition, dehydration, etc. (Fey et al. 2015; Foggin et al. 2023)
  • Other notable diseases
    • Elephant endotheliotropic herpesvirus (abbreviated EEHV) and bovine tuberculosis (caused by the bacterium Mycobacterium tuberculosis) reported in wild populations—but rarely known to kill African elephants (Long et al. 2016; Miller et al. 2019; Miller et al. 2021; Kerr et al. 2023)
      • More research needed to better understand the health impacts on African elephants and potential for disease transmission within/among species (eg, Miller et al. 2019; Kerr et al. 2023; Rob Browning, personal communication, 2025)

Other causes of death (non-comprehensive)

  • Drought and starvation
    • Starvation, rather than dehydration, appears to more commonly kill elephants during drought (Corfield 1973; Shrader et al. 2010; Wato et al. 2016)
      • May have access to water but not nutritious forage
      • Water also plays a role in metabolizing food/stored fat
    • Short-term drought
      • Does not usually harm elephant populations in places where elephants are abundant (Ndlovu et al. 2023)
        • But, can still kill many elephants or make population sizes less stable (Douglas-Hamilton and Wall 2009; Western and Mose 2023)
    • Persistent drought
      • Tends to impact young and very old savanna elephants the most (Dudley et al. 2001; Lee et al. 2016; Ndlovu et al. 2023)
      • Mukeka et al. (2022) report more frequent droughts beginning to cause more elephant deaths than poaching in Kenya
      • Projected to become more frequent in southern Africa, the region with the largest savanna elephant populations (Black et al. 2024)
    • Inability to chew
      • Very worn teeth make it difficult to eat, or very old individuals may no longer have tooth replacements (see Teeth) and die from malnutrition (Eltingham 1991)
  • Injuries
    • Fights between bull males during musth (Hall-Martin 1987)
    • Accidental falls (Douglas-Hamilton and Wall 2009)
      • Calves, especially, may not be able to escape from slippery mud or water wells
    • Landmines (from human warfare) (Eniag et al. 2007)
    • Infections from gunshot wounds or other hunting methods (Poole and Granli 2022)
    • Snares
      • Cause limping, broken bones, joint dislocations, and infections (Poole and Granli 2022)

Family Strides

African elephant calf with family

An African elephant calf walks with family members at the San Diego Zoo Safari Park.

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

 

African elephant calf

African elephant calf

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

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