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

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)

(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

• 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)

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)

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)

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

• See AZA's Species Survival Statistics

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)
      • Also see Orphaned calves
  • 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
  • Humans (Turkalo and Fay 1995)
    • See Threats to Survival

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)

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

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