|An African elephant in South Africa|
Elephants are the largest living land mammals. The largest elephant recorded was one shot in Angola, 1974. It weighed 27,060 pounds (13.5 tons) and stood 13 feet 8 inches tall. Their skin colour is grey.
In the wild, elephants have strong family relationship. Their ways of acting toward other elephants are hard for people to understand. They "talk" to each other with very low sounds. Most elephants sounds are so low, people cannot hear them. But elephants can hear these sounds far away.
- Evolution and extinct relatives
- Anatomy and morphology
- Behaviour and life history
- Conservation issues
- Elephants and humans
Evolution and extinct relatives
Over 161 extinct members of the order Proboscidea have been described. Three major radiations of the proboscideans occurred. The first radiation began with the earliest members, the African Eritherium and Phosphatherium of the late Paleocene. The Eocene included anthracobunids from the Indian subcontinent and Numidotherium, Moeritherium and Barytherium from Africa. These early proboscideans were relatively small and aquatic. Later on, genera such as Phiomia and Palaeomastodon arose; the latter likely inhabited forests and open woodlands. Proboscidean diversity declined during the Oligocene. One notable species of this epoch was Eritreum melakeghebrekristosi of the Horn of Africa, which may have been an ancestor to several later species. The proboscideans diversified at the beginning of the Miocene, with the deinotheres and the mammutids. The former were related to Barytherium and lived in Africa and Eurasia, while the latter may have descended from Eritreum and additionally spread to North America.
The second radiation was represented by the emergence of the gomphotheres in the Miocene, which likely descended from Eritreum and originated in Africa, spreading to every continent except Australia and Antarctica. Members of this group included Gomphotherium and Platybelodon. The third radiation started in the late Miocene and led to the arrival of the elephantids, which descended from, and slowly replaced, the gomphotheres. The African Primelephas gomphotheroides gave rise to Loxodonta, Mammuthus and Elephas. Loxodonta branched off earliest, around the Miocene and Pliocene boundary, while Mammuthus and Elephas came later during the early Pliocene. Loxodonta remained in Africa, while Mammuthus and Elephas spread to Eurasia, and the former reached North America. At the same time, the stegodontids, another proboscidean group descended from gomphotheres, spread throughout Asia, including the Indian subcontinent, China, southeast Asia and Japan. In addition, mammutids continued to evolve into new species, such as the American mastodon.
At the beginning of the Pleistocene, elephantids experienced a high rate of speciation. Loxodonta atlantica became the most common species in northern and southern Africa but was replaced by Elephas iolensis later in the Pleistocene. Only when Elephas became extinct in Africa did Loxodonta become dominant once again, this time in the form of the modern species. Elephas diversified into new species in Asia, such as E. hysudricus and E. platycephus; the latter the likely ancestor of the modern Asian elephant. Mammuthus evolved into several species, including the well-known woolly mammoth. In the Late Pleistocene, most proboscidean species disappeared during the Quaternary glaciation which killed off 50% of genera weighing over 5 kg (11 lb) worldwide.
Proboscideans experienced several evolutionary trends, such as an increase in size, which led to many giant species that stood up to 4 m (13 ft) tall. As with other megaherbivores, including the extinct sauropods, the large size of elephants likely developed to allow them to survive on vegetation with low nutritional value. Their limbs grew longer and the feet shorter and broader. Early proboscideans developed longer mandibles and smaller craniums, while more advanced ones developed shorter mandibles, which shifted the head's centre of gravity. In addition, the skull grew larger, especially the cranium, while the neck shortened to provide better support for the skull. The increase in size lead to the development and elongation of the mobile trunk to provide reach. The number of premolars, incisors and canines decreased. The cheek teeth (molars and premolars) became larger and more specialised. The upper second incisors grew into tusks, which varied in shape from straight, to curved (either upward or downward), to spiralled, depending on the species. Some proboscideans developed tusks from their lower incisiors. Elephants retain certain features from their aquatic ancestry such as their middle ear anatomy and the internal testes of the males.
Several species of proboscideans lived on islands and experienced insular dwarfism. This occurred primarily during the Pleistocene, when some elephant populations became isolated by fluctuating sea levels, although dwarf elephants did exist earlier in the Pliocene. These elephants likely grew smaller on islands due to a lack of large or viable predator populations and limited resources. By contrast, small mammals such as rodents develop gigantism in these conditions. Dwarf proboscideans are known to have lived in Indonesia, the Channel Islands of California, and several islands of the Mediterranean.
Elephas celebensis of Sulawesi is believed to have descended from Elephas planifrons. A dwarf form of Stegodon is known to have existed in Java. Elephas falconeri of Malta and Sicily was only 1 m (3 ft). It was probably a descendant of the straight-tusked elephant. Other descendants of the straight-tusked elephant existed in Cyprus. Dwarf mammoths are known to have lived in Sardinia. Dwarf elephants of uncertain descent lived in Crete, Cyclades and Dodecanese. The Columbian mammoth is known to have colonised the Channel Islands and evolved into the pygmy mammoth. This species reached a height of 1.2–1.8 m (4–6 ft) and weighed 200–2,000 kg (440–4,410 lb). A population of small woolly mammoths survived on Wrangel Island as recently as 4,000 years ago. After their discovery in 1993, they were considered dwarf mammoths. This classification has been re-evaluated and since the Second International Mammoth Conference in 1999, these animals are no longer considered to be true "dwarf mammoths".
Anatomy and morphology
Elephants are the largest extant terrestrial animals. African elephants stand 3–4 m (10–13 ft) and weigh 4,000–7,000 kg (8,800–15,400 lb) while Asian elephants stand 2–3.5 m (7–11 ft) and weigh 3,000–5,000 kg (6,600–11,000 lb). In both cases, males are larger than females. Among African elephants, the forest form is smaller than the savanna form. The skeleton of the elephant is made up of 326–351 bones. The vertebrae are connected by tight joints, which limit the backbone's flexibility. African elephants have 21 pairs of ribs, while Asian elephants have 19–20 pairs.
An elephant's skull is resilient enough to withstand the forces generated by the leverage of the tusks and head-to-head collisions. The back of the skull is flattened and spread out, creating arches that protect the brain in every direction. The skull contains air cavities (sinuses) that reduce the weight of the skull while maintaining overall strength. These cavities give the inside of the skull a honeycomb-like appearance. The cranium is particularly large and provides enough room for the attachment of muscles to support the entire head. The lower jaw is solid and heavy. Because of the size of the head, the neck is relatively short to provide better support. The eye lacks a lacrimal apparatus and relies on the harderian gland to keep it moist. The elephant eye has a durable nictitating membrane to protect the globe. The animal's field of vision is compromised by the location and limited mobility of the eyes. Elephants are considered dichromats and they can see well in dim light but not in bright light. The body temperature of an elephant averages 35.9°C (97°F), similar to a human. Like camels, elephants can raise or lower its temperature a few degrees from the average in response to extreme environmental conditions.
Elephant ears have thick bases with thin tips. The ear flaps, or pinnae, contain numerous blood vessels known as capillaries. To get rid of excess heat from the body, warm blood flows into the capillaries, allowing more heat to be released into the environment. This can occur naturally when the pinnae are still, but the animal can force this effect by flapping them. Larger ear surfaces contain more capillaries, and more heat can be released. Of all the elephants, African bush elephants live in the hottest climates, and thus have the largest ear flaps. Elephants are capable of hearing at low frequencies and are most sensitive at 1 kHz.
The trunk or proboscis, is a fusion of the nose and upper lip, although in early fetal life, the upper lip and trunk are separated. The trunk is elongated and specialised to become the elephant's most important and versatile appendage. It contains up to 150,000 separate muscle fascicles, with no bone and little fat. These paired muscles consist of two major types: superficial (surface) and internal, the former are divided into dorsals, ventrals and laterals, while the latter are divided into transverse and radiating muscles. The muscles of the trunk connect to a bony opening in the skull. The nasal septum is composed of tiny muscle units that stretch horizontally between the nostrils. Cartilage only exists to divide the nostrils at the base. As a muscular hydrostat, the trunk moves by precisely coordinated muscle contractions. The muscles work both with and against each other. A unique proboscis nerve—formed by the maxillary and facial nerves—runs along both sides of the trunk.
Elephant trunks have multiple functions, including breathing, olfaction, touching, grasping and sound production. The animal's sense of smell may be four times as sensitive as that of a bloodhound. The trunk's ability to make powerful twisting and coiling movements allow it to collect food, wrestle with conspecifics, and lift up to 350 kg (770 lb). It can be used for delicate tasks, such as wiping an eye and checking orifices and is capable of cracking a peanut shell without cracking the seed. With its trunk, an elephant can reach items at heights of up to 7 m (23 ft) as well dig up water under mud or sand. It can suck up water both to drink and to spray on itself. An adult Asian elephant is capable of holding 8.5 L (2.2 US gal) of water in its trunk. Elephants will spray dust or grass on themselves, possibly to protect against insects. When underwater, an elephant can use its trunk as a snorkel. Losing the trunk would be detrimental to an elephant's survival, although in rare cases elephants have survived with shortened ones. In addition, one elephant has been observed to graze by kneeling on its front legs, raising on its hind legs and taking in grass with its lips. Elephants may show side preferences when grasping with their trunks: some prefer to twist them to the left, others to the right.
African elephants have two finger-like extensions at the tip of the trunk that allow them to grasp food and bring it to the mouth. Asian elephants have only one, and rely more on wrapping around a food item and squeezing it into their mouths. Asian elephants have more muscle coordination and can perform more complex tasks. Floppy trunk syndrome is a condition of trunk paralysis in African bush elephants. It is caused by the degradation of the peripheral nerves and muscles beginning at the tip.
Elephants usually have 26 teeth: the incisors, known as the tusks, 12 deciduous premolars, and 12 molars. Unlike most mammals, which grow baby teeth and then replace them with a single permanent set of adult teeth, elephants have cycles of tooth rotation throughout their lives. The chewing teeth are replaced six times in a typical elephant's lifetime. Teeth are not replaced by new ones emerging from the jaws vertically as in most mammals. Instead, new teeth grow in at the back of the mouth and move forward to push out the old ones, similar to a conveyor belt. The first chewing tooth on each side of the jaw falls out when the elephant is two to three years old. The second set of chewing teeth falls out when the elephant is four to six years old. The third set is lost at 9–15 years of age, and set four lasts until 18–28 years of age. The fifth set of teeth lasts until the elephant is in its early 40s. The sixth (and usually final) set must last the elephant the rest of its life. Elephant teeth have loop-shaped dental ridges, which are thicker and more diamond-shaped in African elephants.
The tusks of an elephant are modified incisors in the upper jaw. They replace deciduous milk teeth when the animal reaches 6–12 months of age and grow continuously at about 17 cm (7 in) a year. The dentine of the tusk is known as ivory and its cross-section consists of crisscrossing line patterns, known as "engine turning", which create diamond-shaped areas. As a piece of living tissue, a tusk is relatively soft; it is as hard as the mineral calcite. A newly developed tusk has a smooth enamel cap that eventually wears off. Much of the incisor can be seen externally, while the rest is fastened to a socket in the skull. At least one-third of the tusk contains the pulp and some have nerves stretching to the tip. As such it would be difficult to remove it without harming the animal. When removed, ivory begins to dry up and crack if not kept cool and moist. Tusks serve multiple purposes. They are used for digging for water, salt, and roots; debarking or marking trees; and for moving trees and branches when clearing a path. They serve as weapons when fighting, both for offence and defence, and protecting the trunk.
Like humans, who are typically right- or left-handed, elephants are usually right- or left-tusked. The dominant tusk, called the master tusk, is generally more worn down, as it is shorter with a rounder tip. For the African elephants, tusks are present in both males and females, and are around the same length in both sexes, reaching over 3 m (10 ft), but those of males tend to be thicker. In the Asian species, only the males have large tusks. Female Asians have very small ones, or they are absent altogether. Tuskless males do exist and are particularly common among Sri Lankan elephants. Asian males can have tusks as long as Africans, but they are usually much slimmer and lighter; the largest recorded was 3.02 m (10 ft) long and weighed 39 kg (86 lb). The ivory of African forest elephants appear to be the hardest and most elastic. Hunting for elephant ivory in Africa may be leading to natural selection for shorter tusks.
An elephant's skin is generally very tough, at 2.5 cm (1 in) thick on the back and parts of the head. The skin around the mouth, anus and inside of the ear is considerably thinner. Young elephants have brownish or reddish hair, especially on the head and back. Normally the skin of an Asian elephant is covered with more hair than its African counterpart. As elephants age, their hair darkens and becomes sparser, but dense concentrations of hair and bristles remain on the end of the tail as well as the chin, genitals and the areas around the eyes and ear openings. Sparsely spaced hair could allow the animal to lose more heat.
Elephants typically have grey skin, but African elephants look brown or reddish after wallowing in coloured mud. Asian elephants have some patches of depigmentation, particularly on the forehead and ears and the areas around them. An elephant uses mud as a sunscreen, protecting its skin from ultraviolet light. Although tough, an elephant's skin is very sensitive. Without regular mud baths to protect it from burning, insect bites and moisture loss, an elephant's skin suffers serious damage. After bathing, the elephant will usually use its trunk to blow dust onto its body and this dries into a protective crust.
Elephants have difficulty releasing heat through the skin because of their low surface-area-to-volume ratio, which is many times smaller than that of a human. Elephants have even been observed lifting up their legs, presumably in an effort to expose their soles to the air.
Legs, locomotion and posture
To support the animal's weight, an elephant's limbs are positioned more vertically under the body than in most other mammals. The long bones of the limbs have cancellous bone in place of medullary cavities. This strengthens the bones while still allowing blood cell production. Both the front and hind limbs can support an elephant's weight, although 60% is borne by the front. Since the limb bones are placed on top of each other and under the body, an elephant can stand still for long periods of time without using much energy. The circular feet of an elephant have soft tissues or "cushion pads" beneath the manus or pes. These pads help to distribute the weight of the animal. Elephants are incapable of rotating their front limbs, as the ulna and radius are fixed in pronation; the "palm" of the manus faces backward. In addition, the pronator quadratus and the pronator teres are either reduced or absent. As many as five toenails can be found on both the front and hind feet. Elephants appear to have a sesamoid, an extra "toe" similar in placement to a giant panda's extra "thumb", that helps support and distribute their weight.
Elephants can move both forwards and backwards, but cannot trot, jump, or gallop. They use only two gaits when moving on land, the walk and a faster gait similar to running. In walking, the legs act as pendulums, with the hips and shoulders rising and falling while the foot is planted on the ground. With no "aerial phase", the fast gait does not meet all the criteria of running, although the elephant uses its legs much like other running animals, with the hips and shoulders falling and then rising while the feet are on the ground. Fast-moving elephants appear to 'run' with their front legs, but 'walk' with their hind legs and can reach a top speed of 18 km/h (11 mph). At this speed, most other quadrupeds are well into a gallop, even accounting for leg length. Spring-like kinetics could explain the difference between the motion of elephants and other animals.
When moving, the cushion pads expand and contract, and reduce both the pain and noise that would come from a very heavy animal moving. Elephants are capable swimmers. They have been recorded swimming for up to six hours without touching the bottom, and have travelled as far as 48 km (30 mi) at a stretch and at speeds of up to 2.1 km/h (1 mph). While they may doze off while standing, elephants usually sleep lying down.
Behaviour and life history
Ecology and activities
The African bush elephant can be found in habitats as diverse as dry savannas, deserts, marshes, and lake shores, and in elevations from sea level to mountain areas above the snow line. Forest elephants mainly live in equatorial forests, but will enter gallery forests and ecotones between forests and savannas. Asian elephants prefer areas with a mix of grasses, low woody plants and trees, primarily inhabiting dry thorn-scrub forests in southern India and Sri Lanka and evergreen forests in Malaya. Elephants are herbivorous and will eat leaves, twigs, fruit, bark and roots. African elephant are mostly browsers while Asian elephants are mainly grazers. They can consume as much as 150 kg (330 lb) of food and 40 L (11 US gal) of water in a day. Elephants tend to stay near water sources. Major feeding bouts take place in the morning, afternoon and night. At midday, elephants rest under trees and may doze off while standing. Sleeping occurs at night while the animal is lying down. Both males and family groups typically move 10–20 km (6–12 mi) a day, but distances as far as 90–180 km (56–112 mi) have been recorded in the Etosha region of Namibia. They go on seasonal migrations in search of food, water and mates. At Chobe National Park, Botswana, elephant herds travel 325 km (202 mi) to visit the river when the local waterholes dry up.
Because of their large size, elephants have a huge impact on their environments and are considered keystone species. Their habit of uprooting trees and undergrowth can transform savannas into grasslands; when they dig for water during drought, they create waterholes that can be used by other animals. They can enlarge waterholes when they bathe and wallow in them. At Mount Elgon, elephants "excavate" caves that are used by ungulates, hyraxes, bats, birds and insects. Elephants are important seed dispersers; in particular, ingestion and defecation of seeds by African forest elephants tends to have either no effect or a positive effect on their germination and they are typically dispersed in large amounts over great distances. In Asian forests, large seeds require giant herbivores like elephants and rhinoceros for transport and dispersal. This ecological niche cannot be filled by the next largest herbivore, the tapir. Because most of the food they eat goes undigested, their dung can provide food for other animals, such as dung beetles and monkeys. Elephants can have a negative impact on ecosystems. At Murchison Falls National Park in Uganda, the overabundance of elephants has threatened several species of small birds that depend on woodlands. In addition, their weight can compact the soil, which causes the rain to run off, leading to erosion.
The size of adult elephants makes them nearly invulnerable to predators. Calves may be preyed on by lions, spotted hyenas, and wild dogs in Africa and tigers in Asia. The lions of Savuti, Botswana, have adapted to hunting elephants during the dry season, and a pride of 30 lions has been recorded killing individuals between the ages of four and 11 years. Elephants typically coexist peacefully with other herbivores, which will usually stay out their way. Some aggressive interactions between elephants and rhinoceros have been recorded. At Aberdare National Park, Kenya, a rhino attacked an elephant calf and was killed by the other elephants in the group. At Hluhluwe–Umfolozi Game Reserve, South Africa, introduced young orphan elephants went on a killing spree that claimed the lives of 36 rhinos during the 1990s, but ended with the introduction of older males. Elephants tend to have high numbers of parasites, particularly nematodes, when compared to other herbivores. This is due to lower predation pressures that would otherwise kill off many of the individuals with significant parasite loads.
The social lives of male and female elephants are very different. The females spend their entire lives in tight-knit matrilineal family groups, some of which are made up of more than 10 members, including three pairs of mothers with offspring, and are led by the eldest female, or matriarch. She remains leader of the group until death or if she no longer has the energy for the role; a study on zoo elephants showed that when the matriarch died, the levels of faecal corticosterone ('stress hormone') dramatically increased in the surviving elephants. When her tenure is over, the matriarch's eldest daughter takes her place; this occurs even if her sister is present. The older matriarchs tend to be more effective decision-makers.
The social circle of the female elephant does not necessarily end with the small family unit. In the case of African elephants in Amboseli National Park, Kenya, a female's life involves interaction with other families, clans, and subpopulations. Families may associate and bond with each other, forming what are known as bond groups. These are typically made of two family groups. During the dry season, elephant families may cluster together and form another level of social organisation known as the clan. Groups within these clans do not form strong bonds, but they defend their dry-season ranges against other clans. There are typically nine groups in a clan. The Amboseli elephant population is further divided into the "central" and "peripheral" subpopulations.
Some elephant populations in India and Sri Lanka have similar basic social organisations. There appear to be cohesive family units and loose aggregations. In addition, they have been observed to have "nursing units" and "juvenile-care units". In southern India, elephant populations may contain family groups, bond groups and possibly clans. Family groups tend to be small, consisting of one or two adult females and their offspring. A group containing more than two adult females plus offspring is known as a "joint family". Malay elephant populations have even smaller family units, and do not have any social organisation higher than a family or bond group. Groups of African forest elephants typically consist of one adult female with one to three offspring. These groups appear to interact with each other, especially at forest clearings.
The life of the adult male is very different. As he matures, a male begins to spend more time at the edge of his group and associates with outside males or even other families. At Amboseli, young males spend over 80% of their time away from their families when they are 14–15. The adult females of the group start to show aggression towards the male, which encourages him to permanently leave. When males do leave, they either live alone or with other males. The former is typical of bulls in dense forests. Asian males are usually solitary, but do occasionally form groups of two or more individuals; the largest consisted of seven bulls. Larger bull groups consisting of over 10 members occur only among African bush elephants, the largest of which numbered up to 144 individuals. A dominance hierarchy exists among males, whether they range socially or solitarily. Dominance depends on the age, size and sexual condition. Old bulls appear to control the aggression of younger ones and prevent them from forming "gangs". Adult males and females come together for reproduction. Bulls appear to associate with family groups if an oestrous cow is present.
Gestation in elephants typically lasts 18–23 months with interbirth intervals usually lasting four to five years. Births tend to take place during the wet season. Calves are born 85 cm (33 in) tall and weigh around 120 kg (260 lb). The relatively long pregnancy period is maintained by five separate corpus luteums (as opposed to one in most mammals) and gives the foetus more time to develop, particularly the brain and trunk. As such, newborn elephants are precocial and quickly stand and walk to follow their mother and family herd. Typically, only a single young is born, but twins occur on occasion. A new calf is usually the centre of attention for herd members. Adults and most of the other young will gather around the newborn, touching and caressing it with their trunks. For the first few days, the mother is intolerant of other herd members near her young. Alloparenting—where a calf is cared for by someone other than its mother—takes place in some family groups. A mother may have a younger female helper known as an allomother. These allomothers are typically two to 12 years old. When a predator is near, the family group bunches together with the calves in the centre.
For the first few days, the newborn stumbles and falls, and needs the support of its mother. It relies on touch, smell and hearing, as its eyesight is poor. It has little precise control over its trunk, which wiggles around and may cause it to trip. By its second week of life, the calf can walk more firmly and has more control over its trunk. After its first month, a calf can pick up, hold and put objects in its mouth, but cannot suck water through the trunk and must drink directly through the mouth. It is still dependent on its mother and keeps close to her.
For its first three months, a calf relies entirely on milk from its mother for nutrition after which it begins to forage for vegetation and can use its trunk to collect water. At the same time, improvements in lip and leg coordination occur. Calves continue to suckle at the same rate as before until their sixth month, after which they become more independent when feeding. By nine months, mouth, trunk and foot coordination is perfected. After a year, a calf's abilities to groom, drink, and feed itself are fully developed. They still need their mothers for nutrition and protection from predators for at least another year. Suckling bouts tend to last 2–4 min/hr for a calf younger than a year and it continues to suckle until it reaches three years of age or older. Suckling after two years may serve to maintain growth rates, body condition and reproductive ability. Play behaviour in calves differs between the sexes; females run or chase each other, while males play-fight. Females remain in their family groups for life but males leave to live a solitary life or join all-male groups. Elephants have long lifespans, reaching 60–70 years of age. Lin Wang, a captive male Asian elephant, lived for 86 years.
Touching is an important form of communication among elephants. Individuals will greet each other by stroking or wrapping their trunks; the latter also occurs during mild competition. Older elephants use trunk-slaps, kicks and shoves to discipline younger ones. Individuals of any age and sex will touch each other's mouths, temporal glands and genitals, particularly during meetings or when excited. This allows individuals to pick up chemical cues. Touching is especially important for mother–calf communication. When moving, elephant mothers will touch their calves with their trunks or feet when side-by-side or with their tails if the calf is behind them. If a calf wants to rest, it will press against its mother's front legs and when it wants to suckle, it will touch her breast or leg.
Visual displays mostly occur in agonistic situations. Elephants will try to appear more threatening by raising their heads and spreading their ears. They may add to the display by shaking their heads and snapping their ears, as well as throwing dust and vegetation. They are usually bluffing when performing these actions. Excited elephants may additionally raise their trunks. Submissive ones will lower their heads and trunks, as well as flatten their ears against their necks, while those that accept a challenge will position their ears in a V shape.
Elephants produce a number of sounds, usually through the larynx, though some may be modified by the trunk. Perhaps the most well known is the trumpet, which is made during excitement, distress or aggression. Fighting elephants may roar or squeal, and wounded ones may bellow. Rumbles are produced during mild arousal. Some rumbles appear to be infrasonic. Infrasonic calls are important, particularly for long-distance communication, in both Asian and African elephants. For Asian elephants, these calls have a frequency of 14–24 Hz, with sound pressure levels of 85–90 dB and last 10–15 seconds. For African elephants, calls range from 15–35 Hz and can be as loud as 117 dB, allowing communication for many kilometres, with a possible maximum range of around 10 km (6 mi).
At Amboseli, several different infrasonic calls have been identified. A greeting rumble is emitted by members of a family group after having been separated for several hours. Contact calls are soft, unmodulated sounds made by individuals that have been separated from their group and may be responded to with a "contact answer" call that starts out loud, but becomes softer. A "let's go" soft rumble is emitted by a female to signal to the other herd members that it is time to move to another spot. Bulls in musth emit a distinctive, low-frequency pulsated rumble nicknamed the "motorcycle". Musth rumbles may be answered by the "female chorus", a low-frequency, modulated chorus produced by several cows. A loud postcopulatory call may be made by an oestrous cow after mating. When a cow has mated, her family may produce calls of excitement known as the "mating pandemonium".
Elephants are known to communicate with seismics, vibrations produced by impacts on the earth's surface or acoustical waves that travel through it. They appear to rely on their leg and shoulder bones to transmit the signals to the middle ear. When detecting seismic signals, the animals lean forward and put more weight on their larger front feet; this is known as the "freezing behaviour". Elephants posses several adaptations suited for seismic communication. The cushion pads of the feet contain cartilaginous nodes and have similarities to the acoustic fat found in marine mammals like toothed whales and sirenians. A unique sphincter-like muscle around the ear canal constricts the passageway, thereby dampening acoustic signals and allowing the animal to hear more seismic signals. Elephants appear to use seismics for a number of purposes. An elephant running or mock charging can create seismic signals that can be heard at great distances. Seismic waveforms produced by locomotion appear to travel distances of up to 32 km (20 mi) while those from vocalizations travel 16 km (10 mi). When detecting the seismics of an alarm call signalling danger from predators, elephants enter a defensive posture and family groups will pack together.
Intelligence and cognition
Elephants are among the most intelligent species; their psychology is comparable to primates. They exhibit mirror self-recognition, an indication of self-awareness and cognition that has also been demonstrated in some apes and dolphins. One study of a captive female Asian elephant suggested the animals are capable of learning and distinguishing between several visual and some acoustic discrimination pairs. The subject was even able to score a high accuracy rating when retested with the same visual pairs a year later. Elephants are among the few species known to use tools. An Asian elephant was recorded modifying branches and using them as flyswatters. Tool modification by these animals is not as advanced as that done by chimpanzees. Elephants are popularly thought of as having an excellent memory. This could have a factual basis; they possibly have cognitive maps to allow them to remember large-scale spaces over long periods of time. In addition, individuals appear to be able to keep track of the current location of their family members. Scientists debate the extent to which elephants feel emotion. They appear to have some ritual around death and show a keen interest in the bones of their own kind, regardless of whether they are related. A dying or dead elephant seems to elicit attention in others, including those from other groups.
African elephants were listed as vulnerable by the IUCN in 2008, with no independent assessment of the conservation status of the two forms. In 1979, Africa had an estimated minimum population of 1.3 million elephants, with a possible upper limit of 3.0 million. Ten years later in 1989, the population was estimated to be 609,000; with 277,000 in Central Africa, 110,000 in eastern Africa, 204,000 in southern Africa, and 19,000 in western Africa. About 214,000 elephants were estimated to live in the rainforests, less than had previously been thought. From 1977 to 1989, elephant populations declined by 74% in East Africa. Since 1987, losses in elephant numbers accelerated, with savanna populations from Cameroon to Somalia experiencing a decline of 80%. African forest elephants had a total loss of 43%. Population trends in southern Africa were mixed, with anecdotal reports of losses in Zambia, Mozambique and Angola, while populations grew in Botswana and Zimbabwe and were stable in South Africa. Conversely, studies in 2005 and 2007 found populations in eastern and southern Africa were increasing by an average annual rate of 4.0%.
In 2008, the IUCN listed the Asian elephant as endangered due to a 50% population decline over the past 60–75 years. This species once ranged from Syria and Iraq to China (up to the Yellow River) and Java. It is now extinct in these areas, and the current range is heavily fragmented. The total population of Asian elephants is estimated to be around 40,000–50,000, although this may be a loose estimate. It is likely that around half of the population exists in India. Although Asian elephants are declining in numbers overall, particularly in Southeast Asia, the population in the Western Ghats appears to be increasing. CITES lists the species under Appendix I.
The poaching of elephants for their ivory, meat and hides has been one of the major threats to their existence. Historically, numerous cultures made ornaments and other works of art from elephant ivory, and its use rivaled that of gold. Hunting for ivory contributed to the African elephant population decline in the late 20th century. This prompted international bans on ivory imports, starting with the United States in June 1989, and followed by bans in other North American countries, western European countries, and Japan. Around the same time, Kenya destroyed all its ivory stocks. CITES approved an international ban on ivory that went into effect in January 1990.
Following the bans, unemployment rose in India and China, where the ivory industry was important economically. By contrast, Japan and Hong Kong, which were also part of the industry, were able to adapt and were not badly affected. Zimbabwe, Botswana, Namibia, Zambia, and Malawi wanted to continue the ivory trade and were allowed to, since their local populations were healthy, but only if their supplies were from elephants that had been culled or died of natural causes. The ban allowed the African elephant to recover in parts of Africa. In January 2012, hundreds of elephants in Bouba Njida National Park, Cameroon, were killed by Chadian raiders. This has been called "one of the worst concentrated killings" since the ivory ban. Asian elephants are potentially less vulnerable to the ivory trade, as females usually lack tusks. Still, members of the species have been killed for their ivory in some areas, such as Periyar National Park, India.
Other threats to elephants include habitat destruction and fragmentation, which further lead to direct conflicts with humans. The Asian elephant lives in areas with some of the highest human populations. Because they need larger amounts of land than other sympatric terrestrial mammals, they are the first to be affected by human encroachment. In extreme cases, elephants may be confined to small islands of forest among human-dominated landscapes. In addition, elephants cannot coexist with humans in agricultural areas due to their size and food requirements. Elephants commonly trample and consume crops, which contributes to conflicts with humans, and both elephants and humans have died by the hundreds as a result. Mitigating these conflicts is important for conservation.
Elephants and humans
Elephants have been working animals since at least the Indus Valley Civilization and continue to be used in modern times, with 13,000–16,500 working elephants employed in Asia as of 2000. These animals are typically captured from the wild when they are 10–20 years old, when they can be trained quickly and easily, and will have a longer working life. They were traditionally captured with traps and lassos, but since 1950, tranquillisers have been used as they are potentially less harsh. Elephants perform tasks such as hauling loads into remote areas, moving logs into trucks, transporting tourists around national parks, pulling wagons and leading religious processions.
Elephants are valued as working animals over mechanised tools because they can work in relatively deep water, require relatively little maintenance, need only vegetation and water as fuel and can be trained to memorise specific tasks. They can be trained to respond to over 30 commands. Musth bulls can be difficult and dangerous to work with and are chained until the condition passes. Asian elephants are more commonly trained to be working animals, although the practice has also been used in Africa. Leopold II of Belgium is known to have captured and trained elephants in the Belgian Congo during the 19th century. The taming of African elephants continues to the present with the Api Elephant Domestication Centre. In India, many working elephants are alleged to have been subject to abuse. They and other captive elephants are thus protected under the The Prevention of Cruelty to Animals Act of 1960. In northern Thailand, elephants are used to digest coffee beans for Black Ivory coffee.
Elephants were considered formidable instruments of war in antiquity. They were equipped with armour to protect their sides, and their tusks were given sharp points of iron or brass if they were large enough. War elephants were trained to grasp an enemy soldier and toss him to the person riding on them or to pin the soldier to the ground and impale him.
One of the early references to war elephants is in the Indian epic Mahabharata (written in the 4th century BCE, but said to describe events between the 11th and 8th centuries BCE). They were not used as much as horse-drawn chariots by either the Pandavas or Kauravas. During the Magadha Kingdom (which began in the 6th century BCE), elephants began to achieve greater cultural importance than horses, and later Indian kingdoms used war elephants extensively; 3,000 of them were used in the Nandas (5th and 4th centuries BCE) army, while 9,000 may have been used in the Mauryan army (between the 4th and 2nd centuries BCE). The Arthashastra (written around 300 BCE) advised the government to reserve some forests for wild elephants for use in the army, and to execute anyone who killed them. From South Asia, the use of elephants in warfare spread west to Persia and east to Southeast Asia. The Persians used them during the Achaemenid Empire (between the 6th and 4th centuries BCE), while Southeast Asian states first used war elephants possibly as early as the 5th century BCE and continued to the 20th century.
Alexander the Great trained his foot soldiers to injure the animals and cause them to panic during wars with both the Persians and Indians. Ptolemy, who was one of Alexander's generals, used corps of Asian elephants during his reign as the ruler of Egypt (which began in 323 BCE). His son and successor Ptolemy II (who began his rule in 285 BCE) obtained his supply of elephants further south at Nubia. From then on, war elephants were employed in the Mediterranean and North Africa throughout the classical period. Greek king Pyrrhus used elephants in his attempted invasion of Rome in 280 BCE. While they did frighten the Roman horses, they were not decisive and Pyrrhus ultimately lost the battle. The Carthaginian general Hannibal took elephants across the Alps during his war with the Romans and reached the Po Valley in 217 BCE with all of them alive, but they later succumbed to disease.
Zoos and circuses
Elephants were historically kept for display in the menageries of Ancient Egypt, China, Greece and Rome. The Romans in particular pitted elephants against humans and other animals in gladiator events. In the modern era, elephants have traditionally been a major part of zoos and circuses around the world. In circuses, they are be trained to preform numerous tricks. As of 2000, around 1,200 Asian and 700 African elephants are kept in zoos and circuses. The largest captive population is in North America, which has an estimated 370 Asian and 350 African elephants. About 380 Asians and 190 Africans are known to exist in Europe, and Japan has around 70 Asians and 67 Africans. Most African elephants in zoological gardens are bush elephants. Some in Europe, Japan and Africa, though, house forest elephants. These animals do not reproduce well in captivity, due to the difficulty of handling musth bulls and limited understanding of female oestrous cycles. Asian elephants were always more common than their African counterparts in modern zoos and circuses. When CITES listed the Asian elephant under Appendix I in 1975, the number of African elephants in zoos increased in the 1980s, although the import of Asians continued. Subsequently, the US received many of its captive African elephants from Zimbabwe, which had an overabundance of the animals.
Keeping elephants in zoos has been met with some controversy. Proponents of zoos argue that they offer researchers easy access to the animals and provide money and expertise for preserving their natural habitats, as well as safekeeping for the species. Critics claim that the animals in zoos are under physical and mental stress. Elephants have been recorded displaying stereotypical behaviours in the form of swaying back and forth, trunk swaying or route tracing. This has been observed in 54% of individuals in UK zoos. Elephants in European zoos appear to have shorter lifespans than their wild counterparts at only 17 years, although other studies suggest that zoo elephants live as long those in the wild. Supporters of zoos insist that care of animals has improved and activities are used to help reduce stress.
The use of elephants in circuses has also been controversial; the Humane Society of the United States has accused circuses of mistreating and distressing their animals. In testimony to a US federal court in 2009, Ringling Bros. and Barnum & Bailey Circus CEO Kenneth Feld acknowledged that circus elephants are struck behind their ears, under their chins and on their legs with metal-tipped prods, called bull hooks or ankus. Feld stated that these practices are necessary to protect circus workers and acknowledged that an elephant trainer was reprimanded for using an electric shock device, known as a hot shot or electric prod, on an elephant. Despite this, he denied that any of these practices harm elephants. Some trainers have tried to train elephants without the use of physical punishment. Ralph Helfer is known to have relied on gentleness and reward when training his animals, including elephants and lions.
Elephants can experience bouts of aggressive behaviour and engage in actions against humans that have been interpreted as vindictive. In Africa, groups of adolescent elephants damaged homes in villages after cullings in the 1970s and 1980s. In India, male elephants regularly enter villages at night, destroying homes and killing people. Elephants killed around 300 people between 2000 and 2004 in Jharkhand, while in Assam 239 people were reportedly killed between 2001 and 2006. Local people have reported their belief that some elephants were drunk during their attacks, although officials have disputed this explanation. Purportedly drunk elephants attacked an Indian village a second time in December 2002, killing six people, which led to the killing of about 200 elephants by locals.
Elephants have been represented in art since Paleolithic times. Africa in particular contains many rock paintings and engravings of the animals, especially in the Sahara and southern Africa. In the Far East, the animals are depicted as motifs in Hindu and Buddhist shrines and temples. Elephants were often difficult to portray by people with no first-hand experience with them. The ancient Romans, who kept the animals in captivity, depicted anatomically accurate elephants on mosaics in Tunisia and Sicily. At the beginning of Middle Ages, when Europeans had little to no access to the animals, elephants were portrayed more like fantasy creatures. They were often depicted with horse- or bovine-like bodies with trumpet-like trunks and tusks like a boar; some were even given hooves. Elephants were commonly featured in motifs by the stonemasons of the Gothic churches. As more elephants began to be sent to European kings as gifts during the 15th century, depictions of them became more accurate, including one made by Leonardo da Vinci. Despite this, some Europeans continued to portray them in a more stylised fashion. Max Ernst's 1921 surrealist painting The Elephant Celebes depicts an elephant as a silo with a trunk-like hose protruding from it.
Elephants have been the subject of religious beliefs. The Mbuti people believe that the souls of their dead ancestors resided in elephants. Similar ideas existed among other African tribes, who believed that their chiefs would be reincarnated as elephants. During the 10th century AD, the people of Igbo-Ukwu buried their leaders with elephant tusks. The animals' religious importance is only totemic in Africa but is much more significant in Asia. In Sumatra, elephants have been associated with lightning. Likewise in Hinduism, they are linked with thunderstorms as Airavata, the father of all elephants, represents both lightning and rainbows. One of the most important Hindu deities, the elephant-headed Ganesha, is ranked equal with the supreme gods Shiva, Vishnu, and Brahma. Ganesha is associated with writers and merchants and it is believed that he can give people success as well as grant them their desires. In Buddhism, Buddha is said to have been a white elephant reincarnated as a human. In Islamic tradition, the year 570, when the Prophet Muhammad was born, is known as the Year of the Elephant. Elephants were thought to be religious themselves by the Romans, who believed that they worshipped the sun and stars.
Elephants are ubiquitous in Western popular culture as emblems of the exotic, especially since—as with the giraffe, hippopotamus and rhinoceros—there are no similar animals familiar to Western audiences. The use of the elephant as a symbol of the US Republican Party began with an 1874 cartoon by Thomas Nast. As characters, elephants are most common in children's stories, in which they are generally cast as models of exemplary behavior. They are typically surrogates for humans, and their values are ideal human values. Many stories tell of isolated young elephants returning to a close-knit community, such as "The Elephant's Child" from Rudyard Kipling's Just So Stories, Disney's Dumbo and Kathryn and Byron Jackson's The Saggy Baggy Elephant. Other elephant heroes given human qualities include Jean de Brunhoff's Babar, David McKee's Elmer and Dr. Seuss's Horton.
Several cultural references emphasise the elephant's size and exotic uniqueness. For instance, a "white elephant" is a byword for something expensive, useless and bizarre. The expression "elephant in the room" refers to an obvious truth that is ignored or otherwise unaddressed. The story of the blind men and an elephant teaches that reality may be viewed by different perspectives.
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