TAXONOMY & NOMENCLATURE
(Alter 2004) (Fernando et al. 2003)(Gheerbrant et al 1996, 2009) (Nikaido 2003) (Palombo and Villa 2001) (Poulakakis et al 2006) (Rohland et al 2007 (Shoshani 1992,2006 ) (Thenius 1990) (Yang 1996)
Describer: Order established = Illiger (1811). Family describer = Gray
(1821). Revised by Maglio (1973). The Asian Elephant (Elephas) = Linnaeus
(1758).
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Subtribe: Elephantina
Genus: Elephas
Species: Elephas maximus (Asian elephant)
Subspecies: Elephas m. maximus (Sri Lankan Asian elephant)
Subspecies: Elephas m. indicus (Mainland Asian elephant/India and Indochina)
Subspecies: Elephas m. sumatranus
(Sumatran Asian elephant)
Subspecies: Elephas m. borneensis (Borneo Pygmy elephant)
Subtribe: Loxodontina
Genus: Loxodonta
Species: Loxodonta africana
Subspecies: Loxodonta africana africana (bush or savannah
elephant/ eastern, central and southern Africa)
Subspecies: Loxodonta africana cyclotis (forest elephant/central and western Africa)
Taxonomic History and Nomenclature
Subspecies
- E. m. borneensis (Borneo Pygmy Elephants)
- Originally considered distinct subspecies
- Later believed to be feral E. m. sumatranus or E. m. indicus
- Recent genetic studies confirm as a distinct subspecies in Borneo
- Lived on Borneo for up to 300,000 years
Common Name
- Elephant: ele=arch, phant =huge, (Greek).
- Gaja (Sanskrit). Shares root with "garj" meaning "thunder".
- In Hindu mythology water-loving elephants are a harbinger of the monsoon
- Hathi (modern Hindi) from "hastha" or "hasthin" meaning "hand" (alluding to elephant's trunk)
Scientific Name
- Elephas is named for "ele" in Greek, which means arch (referring to the side view profile of the Asian elephant) plus "phant" referring to "fantastic or "huge"
- Maximus comes from "largest" or "greatest" in Latin
Phylogeny
- First elephant-like animals discovered in Moroccan rocks
(Gheerbrant 1996, 2009)
- 55 million year-old Phosphatherium weighed 15 kg (33 lbs)
- 60 million year-old Eritherium weighed 4 to 5 kg (8.8 to 11 lbs)
- Over time, there have been 10 or 11 families in the Order Proboscidea
- Have tusks, modified upper lips and noses,
- Evolutionary trend for size increase from dog-sized to over 4 meters (13 ft.)
- Mastodons diverged from elephant family 24 to 28 million years ago
- Belong to a separate family only distantly related to modern elephants
- Found in Europe, Greece, North America and Central America
- Molecular data: African elephants separate from Asian elephants and mammoths 7.6 million years ago
- New discovery: earliest Asian elephants found in 6.7–5.2 million-year-old rocks in Kenya
- Mammoths (Mammuthus) belong to an extinct genera of the elephant family
- Widespread in Europe, northern Asia, North America and central Mexico, but not South America
- DNA reveals mammoths closer relationship to Asian elephants than to African elephants
- The woolly mammoth (M. primigenius) occupied Europe, British Isles, northern Asia, and as far south as Kansas in the United States
- After arriving in North America, some populations returned to Asia
- The Columbian mammoths (M. columbi) spread throughout North America into Central America
- A dwarf form of the Columbian mammoth survived on the Channel Islands of California
- Extinct about the time of early human contact 12,000 to 13,000 years ago
- Extinct pygmy mammoth skeletons found on Mediterranean island of Crete
and Sardinia
- DNA and morphology shows closer relationship to Mammuthus than to Elephas
-
Modern genera Loxodonta and Elephas both originated in East Africa
- Loxodonta migrated throughout Africa
- Elephas migrated to Asia and Eurasia
- Extinct pygmy elephants on islands of Cyprus and Tilos islands in Mediterranean
- DNA more like modern Elephas
- Not known if pygmy elephants on Sicily and Malta more like mammoths or modern elephants
- Elephants' closest living relatives: hyrax, sea cows, golden moles
- Grouped into proposed new taxon which originated in Africa, Afrotheria, together with aardvarks, elephant shrews, tenrecs
- Close relationships based on molecular evidence, shared anatomy, earth history of plate tectonics
DISTRIBUTION & HABITAT
(Kurt 1990) (Santiapillai & Jackson 1990) (Sukumar 1989, 2008)
Distribution
- Historic range:
- Tigris and Euphrates eastward through Persia
- Indian subcontinent,
- South and Southeast Asia
- China up to the Yangtze-Kiang
- Current populations:
- Indian subcontinent (India, Nepal, Bhutan, Bangladesh)
- Continental Southeast Asia (China, Myanmar, Thailand, Kampuchea, Laos, Vietnam, Malaysia)
- Asian islands (Andaman Islands, Sri Lanka, Sumatra, Borneo)
- Presently occupy only 15% of historic range
- India has 60% of remaining wild populations
- E. m. borneensis' occupied Borneo
in prehistoric times
- not imported from India
- Updated range map available from IUCN (2008).
 |
| Present
distribution and population sizes |
Habitat
- Primary habitat types
- Tropical and subtropical moist broadleaf forests
- Tropical and subtropical dry broadleaf forests
- Move through vegetation zones seasonally; often found along rivers in dry months
- In Southern India: hilly dry scrub through deciduous forest to wet evergreen forest
- Northeastern India and Burma: swampy grasslands on river floodplains; moist deciduous forest on Himalayan slopes
- Sri Lanka: mainly dry deciduous woodland, scrub, grassland, and marshes; small population in rainforest
- India's Andaman Islands: swim across sea between various islands
- Prefer transitional zones between two ecological communities (forests and grasslands)
- Require more access to shade than African elephants
- Inhabit more low visibility habitats compared to African elephants
PHYSICAL CHARACTERISTICS
(Alter 2004)(Fowler & Makata 2006) (Hutchinson et al 2006)(Manoussaki el al 2007) (Mariappa 1986) (Shosani1992)
(Shoshani & Eisenberg 1982)
(Sukumar 1989) (Yokoyama et al 2005)
Body Weight:
- 2000-5500 kg. (4,500-12,000 lbs.)
- Largest living land mammal (several whales are larger)
Body Length:
Tail Length:
- 1.2-1.5 m. (4-5 ft.)
- Tail of Elephas m. boreensis longer than other subspecies - almost touches the ground
Shoulder Height: - Female: 2.24 m. (8 ft.)
- Male: 3.2 m. (10 ft.)
- A bull displayed at the Smithsonian's National Museum of Natural History measured 4 m (13 ft.) at the shoulder
- Shoulder height = forefoot circumference X 2
General
- All elephants have versatile proboscis or trunk, columnar legs, thick skin (pachydermous), and sparse patches of hair
- Smaller than African elephants
- Convex or level back vs more concave in African elephant
- Highest point is at head vs at shoulder in African elephant
- Sri Lankan subspecies is the largest
Skin
- Skin has several textures depending according to location
on body
- Bumpy skin on most of body
- Smooth skin on ears, lips, eyelids, genitalia
- Rough texture isn't found in skin of other mammals
- Generally smoother than the African Elephant’s
- Thick skin - protects against insect bites and weather
- Usually gray. May have white or pink blotches on ears, trunk, head or neck
- Sumatran elephants lightest in color and have least depigmentation (loss of color).
- Sri Lankan elephants darkest and have distinct depigmentation
- Frequently bathe with water, mud or soil to control body temperature
Hair and Nails
- Number of nails don't match number of digits
- Asian elephants: usually 5 nails on forefeet and 4 to 5 nails on hind feet.
- African species: typically 4 to 5 nails on forefeet and 3, 4, or 5 on hind feet
- Infants have "downy" reddish or black hair later replaced by short soft bristles
- Long soft hairs surround ear openings and lower lip
- Short tactile hairs on trunk
- Asians generally harrier than African elephants
Skeleton
- Equals about 15% body weight
- Skull weight equals about 52 kg. (115 lbs.); extensive honeycomb-like spaces reduce skull's weight
- Very short neck brings head close to the center of gravity. Cannot turn head side to side
- Sumatran elephant has 20 sets of ribs, others subspecies have 19 (last 4 are floating)
- African Elephant has 21 sets of ribs
- Side-to-side movement of limbs from the center of the body restricted.
- Increases stability and prevents falls, which can be fatal.
- Feet rest on pads of shock-absorbent, elastic connective tissue, which help support weight
- 34 tail vertebrae
Teeth
- Chewing surfaces of teeth are closed, compressed loops; those of African Elephant are diamond-shaped
- Teeth have a high crown with rasp-like surface, which allows them to chew high fiber materials.
- 26 teeth over lifetime: 2 upper incisors (tusks), 12 deciduous premolars, 12 molars. No canines.
- Six sets of 4 molars molars during lifetime
- Average replacement ages are at 1.5 to 2 years, 6 years, 8 to 10 years, 20 to 25 years and at 50 to 6 0 years.
- The final set is usually lost between 60 and 70 years of age. (Eltringham,1991, p.40).
- Because the teeth change in size and shape throughout life, age of an individual can be estimated
- New molars do not erupt vertically as in most mammals, but grow in from behind, pushing the old worn-out teeth forward and out.
- A single molar can weigh over 5 kg (11 lbs).
Tusks
- Function: dig for water, salt or rocks, debark trees, serve as weapons, protection or rests for the trunk, move branches
- Favors either left or right tusk . One tusk usually shows more wear than the other.
- Elephant incisors develop into
tushes and tusks
- Tushes barely extend past the mouth; replaced by permanent tusks
- Permanent tusks in place by 6 to 12 mos; grow about 17 cm (7 in. )/yr. depending on nutrition
- Large tusks up to 1.8 m (5.9 ft); slimmer and straighter than African's
- Longest tusk on record: 3 meters (119 in.).
- Heaviest weighed 39 kg. (86 lbs)
- Females are tuskless, or have tushes
- Proportion of tuskless males varies widely (0 to 100%). They are called "makhnas."
- Both male and female African elephants usually have tusks
- Tusk microstructure seen in cross section ("Schreger lines") allow identification of various elephant taxa
Trunk
- Technical term is “proboscis," meaning "before the mouth" (Greek)
- Tool for lifting, smelling, spraying dust, grass, and water on body
- Used to transfer water to mouth, not used like a drinking straw
- Also used for sound production, courtship, calf assurance during nursing, behavioral signals and displays
- More "rings" (annulations) than in African Elephant's and perhaps is more extendable
- Formed by a fusion of the nose and upper lip
- One prehensile “finger” at tip of trunk (mammoths had one also); African elephant has two "fingers"
- Three types of tactile receptors at tip; most sensitive part of the trunk
- Contains as many as 150,000 muscle units
- Muscle masses aligned in radial, longitudinal, and oblique layers for extraordinary flexibility
- Tongues of vertebrates, arms of octopus and tentacles of squid have similar flexibility
- 70% of air inhaled is through the trunk, the rest comes through the mouth.
- Uses trunk to transfer water to mouth, not used for drinking;can hold up to 5 liters (1.3 gal) water
- Uses trunk to transfer chemical pheromones to receptors in the mouth
- Trunk tip's contain nerve endings with heightened sensitivity to vibrations
- When placing trunk tip on ground, can detect vibrations of running animals
Ears
- Smaller than the African elephant's.
- E.m. borneensis has larger outer ear (pinna) than other subspecies
- Thermo regulation - positive correlation between the number of times an elephant flaps its ears and air temperature.
- Cochlea's curved structure may facilitate sensitivity to low frequencies.
- Can hear approximately 8Hz. – 12 kHz
- For comparison: humans hear up to 19kHz and dogs to 44 kHz
- Represent best sensitivity of all mammals so far tested (Langbauer 2000)
- May be able to detect seismic vibrations from thunderstorms, animal hooves through feet
Glands
- Two pectoral mammary glands produce milk that is 83.82% water, 11.82% albuminoids and sugar, 3.89% fat and 0.47% ash or mineral matter.
- Cheek (temporal) glands located midway between eyes and ears. Drain during times of excitement such as fighting, mating or in musth
(once or twice a year)
- Produce pheromones that stimulate other elephants' behavior
- Present in both males and females; much more active in males
- Tear ducts are vestigial. Herderian glands lubricate the eyes.
Internal Organs
- Weight of heart 19 kg (42 lbs); about 0.5 % of body weight. Unique shape like the African Elephant with bifed apex and paired venae cavae
- Heart rate 28 beats per minute standing; 35 lying down
- Rate of respiration 4-12 per minute
- Brain weight 5.5 kg (12 lbs). Large temporal lobes indicate heightened memory capabilities. Larger than humans', relative to body size
- Absence of gall bladder (have low-fat diet)
- Pancreas does not come in contact with spleen, unlike most animals.
- Testes located near kidneys inside abdomen; heat-sensitive sperm ok because of elephant's lower body temperature
- Lungs attached to diaphragm; typically breathe using diaphragm rather than expansion of rib cage
Eyesight
- Eye is small in relation to body size
- Ability to see color - probably similar to humans who are color blind
- Total visual field is a sweep of 313° out of 360° with a 47° blind spot (compared to 357° visual field for a horse)
- No true tear ducts
- Good in dull light, considerably reduced in bright light
Sexual Dimorphism
- Males have large trunk bases, bulges below and in front of eyes, and swelling above the eyes. Females have narrower trunk bases and lack prominent bulge above eyes
- Male back is more convex and curves more gradually into hindquarters; female's is straighter and "boxier" with vertical hindquarters.
- Males considerably larger than females of same ages.
Other Physical Characteristics
-
Folds in brain resemble that of humans and porpoises
- Highly sensitive Jacobson's organ on roof of mouth used for detecting sex pheromones
BEHAVIOR & ECOLOGY
(Eltringham 1991) (Fernando and Lande 2000) (Hart et al 2001) (McKay 1973) (Moss 1990) (O'Connell-Rodwell 2001)
(Payne & Langbauer 1992)(Poole 1996) (Pringle 2008) (Rasmussen and Krishnamurthy 2000) (Santipelli &Suprahman 1986) (Schulte 2006)(Shoshani & Eisenberg 1982) (Sukumar 1989)
Activity Cycle
- In a study of Asian Elephants at Lahugala Tank, males were recorded as having the following activity cycle:
91.1% = Feeding
5.4% = Walking
1.4% = Resting
1.8% = Bathing
0.1% = Drinking
0.2% = All other activities
- Another study at Lahagula Tank records the following activity cycle of solitary males:
93.5% = Feeding
1.9% = Walking
0.3% = Contact promoting
3.3% = Play/Aggression
1.0% = All other activities
- Female elephant herds spend 70-90% of time feeding.
- Male activity shows one peak at around 8 am and another between 4-5 pm. Female herds peak around 10 am and then later between 3:30 and 6 pm.
- In a study at Ceylon's Wilpattu National Park
for
individuals described as secretive and shy
- Feeding is intermittent during 24 hour period
- Drinking is typically early evening and just before daybreak
- Long distance travel usually accomplished at night
Territory Size
- No evidence of territoriality
- Home range size usually small, similar to African Elephants (approx. 14-52 sq.km)
- Shifts locale with wet/dry season
- Availability of resources (water, food, mates) and human presence all influence home range size
- Female groups may overlap and coordinate migrations
Social Groups
- Highly social animals with complex behavior
- A matriarch, typically the eldest female, heads a related group of females.
- Study of female groups's genetics at Ruhuna National Park in Sri Lanka show all individuals related to a single female
- Herds consist of about 8-12 individuals, but sizes can vary. (African Elephant groups: 10 to 20 individuals)
- Life centers around the calves in both African and Asian elephant societies.
- Females share equally in the care of the young.
- Teenage males leave natal herd & often form bachelor groups
- Adult male typically described as solitary; no close bonds with other independent males; spend time in families only when following females in estrus
- New studies show the males form loose associations
Hierarchy
- Males in musth are dominant to nonmusth males
- Teenage Asian elephants (15-20 years) go through a juvenile (moda) musth in which they secrete a sweet odor
- Scent signals subordinate status, eliminating unnecessary conflict with older, dominant males.
Aggression
- Bulls in musth once or twice a year may display aggressive behavior
- Having a male in musth decreases aggressive behavior of other males
- Aggressive behaviors include: spreading ears; raising head with jaw "tucked in", mock charging, abrupt head shake which makes ears flap, throwing objects in direction of opponent
- Elephant charge: up to 40 km/hr (25mph)
Play
- Male calves are more likely to leave mothers to play; engage in head sparring, mounting, charging, shoving, and chasing
- Female calves play chasing and running games, throw sticks, and may “attack” imaginary enemies
- Both sexes play rough and tumble games, clamber onto each other
Communication
Vocal Cues
- Large repertoire of vocalizations, including:
- Low frequency, long distance calls to maintain contact between roving individuals or groups
- Close range, high pitched calls indicate mood
- Loud trumpeting indicates alarm, surprise
- Low snorts signal changes in immediate environment, alerting herd
Tactile Cues
- Family members often touch while standing; may rub with a foot or slap with the trunk
- Trunks are used in greeting: A lower-ranking animal will insert its trunk tip into the other’s mouth
- Trunk may be held out to an approaching elephant as a greeting
- Trunk is also used in caressing, twining, wrestling, and checking reproductive status
- Mothers may guide their calf by gripping its tail
Visual Cues
- Signal for hormonal state:
- Bull's musth walk - head erect, ears wide, ear waves of one ear, a low pulsing growl
- Trunk curling and uncurling
- Urine dribble
- Cow's estrous walk - looking back over shoulder as walk away
- Signals for apprehension/submission
- Jaw out, touching one's own temporal gland or face
- Trunk twitching back and forth
- Swaying side to side
- Backing into side of more dominant animal
Olfaction/Scent Marking
- Rely heavily on long lasting chemical cues, which travel over short or long distances
- Lift trunks to detect wind borne scents for first clues to sources of danger
- Survival depends on reading scents of landscapes, pathways, mineral and salt sources, waterholes
- Even after long separations, chemical cues help reestablish kin and friendship bonds
- Sniff breath, mouths, temporal glands, genitals, urine and dung to determine emotional and physiological states of others
- Urine alone contains several thousand chemical compounds packed with messages
- Sex pheromones allow determination of fitness and location of opposite sex
Other senses
- Information from environment via low frequency ground vibrations may be first detected by elephant's feet and trunk tip
- Distant thunderstorms, footsteps of running animals, vocalizations, that travel through ground
Locomotion
- Move with one basic gait, known as the rack or pace (three feet are on the ground most of the time); one foot always on ground
- Normal walking speed 2.5-3.7 mph (4-6 km/hr). Charging speed can reach 15.2 mph (6.8 meters/sec) (Hutchinson 2006)
- Move very slowly while feeding, or walk quickly. Juveniles may run when playing, but adults only run in flight or attack.
- Swim readily at rate of 1.3 mph (2 km/hr). Can stay afloat for up to 6 hours and cover distances of 30 miles (48 km) at a stretch
- Can not jump; even a shallow ditch poses a barrier
- Can not trot, canter, or gallop
Tool Use
- May use branches and plant fronds to swat flies, dislodge parasites, scratch an itch
- Known to drop boulders on fences to destroy them or to cut off electricity
Interspecies Interaction
- Do not appear frightened by other animals; usually ignore them
- Confrontations with Indian rhinoceros probably very rare in wild, but have occurred with domestic elephants that have been ridden into rhino's territory
- Mutualistic relationship with some species of birds, such as egrets and piapiac.
- Birds use elephants as a vantage point to spot prey; pick off ticks or lice attached to the elephant.
- Host several parasites, including louse, warble flies, mosquitoes and leeches
- Prodigious amounts of dung disperses many seeds and helps enrich soils
- Dung beetles and termites carry dung underground
- Like the African Elephant, function in role of ecosystem "engineer"
and as a keystone species
- Maintain trails and open access to water for other species
- Create microhabitats by shredding trees for small vertebrate species such as lizards
- Uproot small trees (which would invade open grassland areas, shifting habitat to new forests)
- Enhances habitat for other grazing animals and their predators
DIET & FEEDING
(Poole 1996)
(Santiapelli & Suprahman 1986)
(Sukumar 1989, 1994)
Anatomy and Physiology
- Trunk and flexible "finger" at tip allows precision in feeding; can choose a single flower
- Molar teeth
rasp-like with ridges of enamel suited for grinding coarse and silica-rich plants
- Considered a non-ruminant but has large caecum and colon for
fermentation and digestion
- Digestive strategy: pass large amounts of low quality forage pass through digestive tract quickly
- Have no gall bladder; eat low-fat, high fiber diet
- Defecate as often as 15 to 20 times per day; wild herd members often defecate at the same time
Food Items and Feeding Strategies
- Are both browsers (feeding on shrubs and trees) and grazers (feeding on grass)
- Can consume many species of plants, as well as twigs and bark but usually only a few plant taxa
- 112 species of plants in a study in southern India, 400 species recorded in Malaysia
- Choice of plants varies with seasons
- Feed on mostly browse plants during dry season
- Shift to grass in first part of wet season;
- Return to browse feeding after heavy rains (when grass less nutritious)
- Strategies for obtaining grass
include:
- Dislodge clumps with feet, gather a pile of grass, grab with trunk, smack against leg to knock off dirt, put in mouth when dirt-free
- Wash clumps by moving back and forth with trunk in water
- Usually do not feed for more than a few days in one place
- Drink every day when available; stay close to a source of fresh water
- Can ingest over 100 liters (26 gallons) at one time
- Drink up to 225 liters (59 gallons) per day
- If no water available, dig with forefeet and trunks for subsoil water; wallows created in this manner
- Anecdotal evidence reports elephants fond of alcohol; reports of raiding illicit liquor stores in the forest (Sukamar 89)
- Adults eat approx. 150kg (330 lbs) daily; approximately 1.5 – 2% (dry weight) of body weight
- Salt needed in diet; migrate to deposits regularly and excavate with tusks if necessary
REPRODUCTION & DEVELOPMENT
(Allen 2006) (Alter 2004)(Langbauer 2000) (Moss 1990)
(Shoshani & Eisenberg 1982)(Sukumar 1994)
Courtship
- Female urine contains a sex pheromone that increases before ovulation (African Elephant's pheromone differs)
- Female emits loud infrasonic call when in estrus to attract males
- Important because receptive time may last one day; calls persist for 5 days
- Audible up to 4 km (2.4 miles) away
- Bulls sniff female genitals and urine, place trunk tip into mouth:
- Jacobson's organ on roof of mouth detects presence of the sex pheromone
- Prior to copulation, wrestle with intertwined trunks, bite each other’s necks, male attempts mounting
- Females exercise some choice in mates by running away and/or not standing still
- Females in estrus prefer breeding with males in musth
Reproduction
- Mounts last about 30 seconds
- Mating and births occur any time during year; may peak during heavy rains in some regions
- In captivity, estrus cycle lasts 13 to 14 weeks (if female is not pregnant)
- Females who have conceived or with young may have 4 - 5 years between estrus cycles
- Only one in five follicular cycles, each lasting 2-3 weeks, produce a fertile egg
- Males can breed at any time, not only when in musth (Hindu for "intoxicated")
- Musth levels of testosterone in male up to 20 times higher than normal
- Musth lasts about 4 months in wild males
- Females prefer musth males
Reproductive Rate:
- Most females give birth for the first time at 16-17 years.
- In optimal habitats, calving intervals are 2.5-4 years. In less favorable conditions, calving intervals are 5-8 years.
- Cows can calve past age 50. Two captive elephants (Meennakshi and Tara) calved at 54 and 62 years, respectively.
Gestation
- Lasts 18-22 months, the longest gestation period of any mammal
Life Stages
- Birth
- Litter size: Usually one calf; rarely twins (structure of uterus allows births several weeks apart)
- Weight: 75-115kg (165-254 lbs)
- Height: 75-100 cm (2.5-3.5 ft)
- Calves are very hairy compared to adults. Hairier than African elephant calves.
- Infant (< 1 year old)
- Infants may have many female juvenile helpers - allomothers or "aunties" (great survival advantage)
- Helpers allow mother to feed and rest; important for lactation
- Can stand on feet shortly after birth
- Follow the mother in her daily routines within a few days
- During first three months develop motor skills
- Females attracted to young calves
- Juvenile
- Females are attracted to young calves; often minister to their needs so mothers can spend more time feeding and resting, which are important for lactation
- Age of weaning 18 months to 3 years or longer;
- May be forced to wean when younger sibling is born.
- Around age of weaning, differences in sexes begin to appear
- Females strengthen bonds with family
- Males become more independent, leaving family for longer periods to feed, explore or play
- Initial period of rapid growth slows at about 5 years; decline in growth rate is greater for females than for males
- Females care for young of both sexes during 10-15 years of development until sexual maturity
- Subadult
- Can be difficult to distinguish from juveniles
- No fixed age of puberty; may depend on available nutrition
- Males often seen away from herds
- Penis large, although subadult males are not reproductively active
- Females may exhibit swelling of mammae
- Adult
- Age of sexual maturity varies depending on nutrition
- Bulls 14-15 (as early as 9) years
- Cows 8-13 years
- Bulls may not mate until late teens or twenties due to social hierarchy
- Males are distinguishable by enlargement of head and distinct penis sheath; no testicles visible
- Females and males often hard to distinguish because female's clitoris is large
- Females have square backs
- Males continue to grow at continuous slow rate. Female growth plateaus between 15 and 30 years
- Males in wild enter musth around age 30
Longevity
- Comparable to human longevity
- May live 60-70 years in the wild. Over 80 years in captivity
Mortality