Elephants are large mammals of the family Elephantidae and the order Proboscidea. Three species are recognised, the African bush elephant (Loxodonta africana), the African forest elephant (L. cyclotis), and the Asian elephant (Elephas maximus). Elephants are scattered throughout sub-Saharan Africa, South Asia, and Southeast Asia. Elephantidae is the only surviving family of the order Proboscidea; other, now extinct, members of the order include deinotheres, gomphotheres, mammoths, and mastodons. Male African elephants are the largest extant terrestrial animals and can reach a height of 4 m (13 ft) and weigh 7,000 kg (15,000 lb). All elephants have several distinctive features, the most notable of which is a long trunk or proboscis, used for many purposes, particularly breathing, lifting water, and grasping objects. Their incisors grow into tusks, which can serve as weapons and as tools for moving objects and digging. Elephants' large ear flaps help to control their body temperature. Their pillar-like legs can carry their great weight. African elephants have larger ears and concave backs while Asian elephants have smaller ears and convex or level backs.
Elephants are herbivorous and can be found in different
habitats including savannahs, forests, deserts, and marshes.
They prefer to stay near water. They are considered to be keystone species due to their impact on their environments. Other
animals tend to keep their distance from elephants while predators, such
as lions, tigers, hyenas,
and any wild dogs, usually target only young elephants (or
"calves"). Females ("cows") tend to live in family groups,
which can consist of one female with her calves or several related females with
offspring. The groups are led by an individual known as the matriarch, often the oldest cow. Elephants have a fission–fusion
society in which multiple family groups
come together to socialise. Males ("bulls") leave their family groups
when they reach puberty and may live alone or with other males. Adult bulls
mostly interact with family groups when looking for a mate and enter a state of
increased testosterone and aggression known as musth,
which helps them gain dominance and reproductive success. Calves are the centre of
attention in their family groups and rely on their mothers for as long as three
years. Elephants can live up to 70 years in the wild. They communicate by
touch, sight, smell, and sound; elephants use infrasound, and seismic
communication over long distances.
Elephant intelligence has been compared with that of primatesand cetaceans. They appear to have self-awareness and show empathy for dying or dead individuals of their kind.
African elephants are listed as vulnerable by the International
Union for Conservation of Nature (IUCN)
while the Asian elephant is classed as endangered. One of the biggest threats to elephant populations is
the ivory trade, as the animals are poached for their ivory tusks. Other threats to wild
elephants include habitat destruction and conflicts with local people. Elephants are used
as working
animals in Asia. In the past, they were
used in war; today, they are often controversially put on display in zoos, or
exploited for entertainment in circuses. Elephants are highly recognisable and
have been featured in art, folklore, religion, literature, and popular culture.
Etymology
The word "elephant" is based on
the Latin elephas (genitive elephantis) ("elephant"), which is the Latinised form of the Greek ἐλέφας (elephas) (genitive ἐλέφαντος (elephantos)), probably from a non-Indo-European language, likely Phoenician. It is attested in Mycenaean
Greek as e-re-pa (genitive e-re-pa-to) in Linear B syllabic script. As in Mycenaean
Greek, Homer used
the Greek word to mean ivory, but after
the time of Herodotus, it also
referred to the animal. The word "elephant" appears in Middle
English as olyfaunt(c.1300) and was borrowed from Old French oliphant (12th century). Loxodonta,
the generic name for the African elephants, is Greek for "oblique-sided
tooth".
Taxonomy
Classification, species and subspecies
Elephants belong to the family Elephantidae, the sole remaining family within the order Proboscidea which belongs to the superorder Afrotheria. Their closest extant relatives are the sirenians (dugongs and manatees) and the hyraxes,
with which they share the clade Paenungulata within the superorder Afrotheria. Elephants and sirenians are further grouped in the
clade Tethytheria. Three species of elephants are recognised;
the African
bush elephant (Loxodonta africana) and forest elephant (Loxodonta
cyclotis) of sub-Saharan Africa, and the Asian elephant (Elephas maximus) of South and Southeast Asia. African elephants have larger ears, a concave back,
more wrinkled skin, a sloping abdomen, and two finger-like extensions at the
tip of the trunk. Asian elephants have smaller ears, a convex or level back,
smoother skin, a horizontal abdomen that occasionally sags in the middle and
one extension at the tip of the trunk. The looped ridges on the molars are narrower in the Asian elephant while those of
the African are more diamond-shaped. The Asian elephant also has dorsal bumps
on its head and some patches of depigmentation on its skin. In general, African elephants are
larger than their Asian cousins.
Swedish zoologist Carl Linnaeus first described the genus Elephas and an elephant
from Sri Lanka (then known as Ceylon) under the binomialElephas maximus in 1758. In 1798, Georges Cuvier classified the Indian elephant under the binomial Elephas indicus. Dutch zoologist Coenraad Jacob
Temminck described the Sumatran elephant in 1847 under the binomial Elephas sumatranus. English zoologist Frederick Nutter
Chasen classified all three as subspecies of the Asian elephant in 1940.[10] Asian
elephants vary geographically in their colour and amount of depigmentation.
The Sri
Lankan elephant (Elephas maximus maximus) inhabits Sri Lanka, the Indian elephant (E. m. indicus) is native to mainland Asia (on the Indian subcontinent and Indochina), and the Sumatran elephant (E. m. sumatranus) is found in Sumatra. One disputed subspecies, the Borneo elephant, lives in northern Borneo and
is smaller than all the other subspecies. It has larger ears, a longer tail,
and straighter tusks than the typical elephant. Sri Lankan zoologist Paules
Edward Pieris Deraniyagala described it in
1950 under the trinomial Elephas
maximus borneensis, taking as his type an illustration in National Geographic. It was subsequently subsumed under either E. m. indicus or E. m. sumatranus.
Results of a 2003 genetic analysis indicate its ancestors separated from the mainland population about
300,000 years ago. A 2008 study found that Borneo elephants are not
indigenous to the island but were brought there before 1521 by the Sultan of Sulu from Java,
where elephants are now extinct.
The African elephant was first named by German naturalist Johann
Friedrich Blumenbach in 1797 as Elephas africana. The genus Loxodonta was
commonly believed to have been named by Georges Cuvier in 1825. Cuvier spelled
it Loxodonte and an anonymous author romanised the spelling to Loxodonta; the International
Code of Zoological Nomenclature recognises
this as the proper authority. In 1942, 18 subspecies of African elephant
were recognised by Henry Fairfield
Osborn, but further morphological data has
reduced the number of classified subspecies, and by the 1990s, only two
were recognised, the savannah or bush elephant (L. a. africana)
and the forest elephant (L. a. cyclotis); the latter has smaller and more rounded ears and
thinner and straighter tusks, and is limited to the forested areas of western and Central Africa. A 2000 study argued for the elevation of the two
forms into separate species (L. africana and L. cyclotisrespectively)
based on differences in skull morphology. DNA studies published in 2001
and 2007 also suggested they were distinct species while studies in 2002
and 2005 concluded that they were the same species. Further studies (2010,
2011, 2015) have supported African savannah and forest elephants' status as
separate species. The two species are believed to have diverged 6 million
years ago and in 2016, DNA sequence analysis showed that L. cyclotis is more closely related to the extinct Palaeoloxodon antiquus, than it is to L. africana. Some
evidence suggests that elephants of western Africa are a separate
species, although this is disputed. The pygmy elephants of the Congo Basin, which have been suggested to be a separate species (Loxodonta pumilio) are probably forest elephants whose small size and/or
early maturity are due to environmental conditions.
Evolution and extinct relatives
Over 161 extinct members and three major evolutionary radiations of the order Proboscidea have been recorded. The
earliest proboscids, the African Eritherium and Phosphatherium of the
late Paleocene, heralded the first radiation. The Eocene included Numidotherium, Moeritherium, and Barytherium from Africa. These animals 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 beginning of the Miocene saw the second diversification, with the appearance of 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 spread to
North America.
The second radiation was represented by the emergence of
the gomphotheres in the Miocene, which likely evolved
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 diverged
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. 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. The Pleistocene also saw the arrival of Palaeoloxodon namadicus, the largest terrestrial mammal of all time. 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 disappeared from 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 vanished 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
5 m (16 ft) tall. As with other megaherbivores, including the extinct sauropod dinosaurs, 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. The feet were originally plantigrade and developed into a digitigrade stance with cushion pads and the sesamoid bone providing support. Early proboscideans
developed longer mandibles and smaller craniumswhile
more derived ones developed shorter mandibles, which shifted the
head's centre
of gravity. The skull grew larger, especially
the cranium, while the neck shortened to provide better support for the skull.
The increase in size led to the development and elongation of the mobile trunk
to provide reach. The number of premolars, incisors and caninesdecreased. The cheek teeth (molars and premolars)
became larger and more specialized, especially after elephants started to
switch from C3-plants to C4-grasses, which caused their teeth to undergo a three-fold
increase in teeth height as well as substantial multiplication of lamellae
after about five million years ago. Only in the last million years or so did
they return to a diet mainly consisting of C3 trees and shrubs. 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 incisors. Elephants retain
certain features from their aquatic ancestry, such as their middle ear anatomy and the internal testes of the males.
There has been some debate over the relationship of Mammuthus to Loxodonta or Elephas. Some DNA studies
suggest Mammuthus is more closely related to the former while
others point to the latter. However, analysis of the complete mitochondrial genome profile of the woolly mammoth (sequenced in 2005)
supports Mammuthus being more closely related to Elephas. Morphological evidence supports Mammuthus and Elephas as sister taxa while comparisons of protein albumin and collagen have concluded that all three genera are equally
related to each other. Some scientists believe a cloned mammoth embryo could
one day be implanted in an Asian elephant's womb.
Dwarf species
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. Elephas falconeri of Malta and Sicily was
only 1 m (3 ft) and had probably evolved from the straight-tusked
elephant. Other descendants of the
straight-tusked elephant existed in Cyprus.
Dwarf elephants of uncertain descent lived in Crete, Cyclades, and Dodecanese while dwarf mammoths are known to have lived
in Sardinia. The Columbian mammoth 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, now 140 km (87 mi) north of the Siberian
coast, 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 living 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
savannah 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 or 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 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. Lacking a lacrimal apparatus, the eye relies on the harderian glandto keep it moist. A durable nictitating membrane protects the eye 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 core body temperature averages 35.9 °C
(97 °F), similar to a human. Like all mammals, an elephant can raise or
lower its temperature a few degrees from the average in response to extreme
environmental conditions.
Ears
Elephant ears have thick bases with thin tips. The ear
flaps, or pinnae, contain numerous blood vessels called capillaries. Warm blood flows into the capillaries, helping to
release excess body heat into the environment. This occurs when the pinnae are
still, and the animal can enhance the 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 have the
largest ear flaps. Elephants are capable of hearing at low frequencies and
are most sensitive at 1 kHz.
Trunk
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 divides 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 allows it to collect food, wrestle with other
elephants, and lift up to 350 kg (770 lb). It can be used
for delicate tasks, such as wiping an eye and checking an orifice, and is
capable of cracking a peanut shell without breaking the seed. With its
trunk, an elephant can reach items at heights of up to 7 m (23 ft)
and dig for water under mud or sand. Individuals may show lateral preference
when grasping with their trunks: some prefer to twist them to the left, others
to the right Elephants can suck up water both to drink and to spray on
their bodies. An adult Asian elephant is capable of holding 8.5 L
(2.2 US gal) of water in its trunk. They will also spray dust or
grass on themselves. When underwater, the elephant uses its trunk as a snorkel.
The African elephant has two finger-like extensions at
the tip of the trunk that allow it to grasp and bring food to its mouth. The
Asian elephant has only one, and relies more on wrapping around a food item and
squeezing it into its mouth. Asian elephants have more muscle coordination
and can perform more complex tasks. Losing the trunk would be detrimental
to an elephant's survival, although in rare cases, individuals have
survived with shortened ones. 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. Floppy trunk
syndrome is a condition of trunk paralysis in African bush elephants caused by the degradation
of the peripheral
nerves and muscles beginning at the tip.
Teeth
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 are polyphyodonts that 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. 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.
Tusks
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. A
newly developed tusk has a smooth enamel cap that eventually wears off. The dentine 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. 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 pulpand some have nerves stretching to the tip. Thus 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. When
fighting, they are used to attack and defend, and to protect 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
up to 3 m (10 ft), but those of males tend to be
thicker. In earlier times, elephant tusks weighing over 200 pounds (more
than 90 kg) were not uncommon, though it is rare today to see any over 100
pounds (45 kg).
In the Asian species, only the males have large tusks.
Female Asians have very small ones, or none at all. Tuskless males exist
and are particularly common among Sri Lankan elephants. Asian males can
have tusks as long as Africans', but they are usually slimmer and lighter; the
largest recorded was 3.02 m (10 ft) long and weighed 39 kg
(86 lb). Hunting for elephant ivory in Africa and Asia has led
to natural
selection for shorter tusks and
tusklessness.
Skin
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.
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. Calves have brownish or reddish hair, especially on the head and back. As
elephants mature, 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.
Normally the skin of an Asian elephant is covered with more hair than its
African counterpart.
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. They 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 haematopoiesis. 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. Elephants are incapable of
rotating their front legs, as the ulna and radius are fixed in pronation; the "palm" of the manus faces
backward. The pronator
quadratusand the pronator teres are either reduced or absent. The circular
feet of an elephant have soft tissues or "cushion pads" beneath
the manus or pes, which distribute the weight of the animal. They
appear to have a sesamoid, an extra "toe" similar in placement to
a giant panda's extra "thumb", that also helps in weight
distribution. As many as five toenails can be found on both the front and
hind feet.
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. During locomotion, 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).
Internal and sexual organs
The brain of an elephant weighs 4.5–5.5 kg
(10–12 lb) compared to 1.6 kg (4 lb) for a human brain. While
the elephant brain is larger overall, it is proportionally smaller. At birth,
an elephant's brain already weighs 30–40% of its adult weight. The cerebrum and cerebellum are well developed, and the temporal lobes are so large that they bulge out
laterally. The throat of an elephant appears to contain a pouch where it
can store water for later use.
The heart of an elephant weighs 12–21 kg
(26–46 lb). It has a double-pointed apex, an unusual trait among mammals. In addition,
the ventricles separate near the top of the heart, a trait they
share with sirenians. When standing, the elephant's heart beats
approximately 30 times per minute. Unlike many other animals, the heart rate
speeds up by 8 to 10 beats per minute when the elephant is lying down. The
blood vessels in most of the body are wide and thick and can withstand high
blood pressures. The lungs are attached to the diaphragm, and breathing relies mainly on the diaphragm rather
than the expansion of the ribcage. Connective tissue exists in place of the pleural cavity. This may allow the animal to deal with the pressure
differences when its body is underwater and its trunk is breaking the surface
for air, although this explanation has been questioned. Another
possible function for this adaptation is that it helps the animal suck up water
through the trunk. Elephants inhale mostly through the trunk, although
some air goes through the mouth. They have a hindgut fermentation system, and their large and small intestines
together reach 35 m (115 ft) in length. The majority of an elephant's
food intake goes undigested despite the process lasting up to a day.
A male elephant's testes are located internally near the
kidneys. The elephant's penis can
reach a length of 100 cm (39 in) and a diameter of 16 cm
(6 in) at the base. It is S-shaped when fully erect and has a
Y-shaped orifice. The female has a well-developed clitoris at up to 40 cm (16 in). The vulva is
located between the hind legs instead of near the tail as in most mammals.
Determining pregnancy status can be difficult due to the animal's large abdominal cavity. The female's mammary glands occupy the space between the front legs, which puts
the suckling calf within reach of the female's trunk. Elephants have a
unique organ, the temporal
gland, located in both sides of the head. This
organ is associated with sexual behaviour, and males secrete a fluid from it
when in musth. Females have also been observed with secretions
from the temporal glands.
Behaviour
and life history
Ecology and activities
The African bush elephant can be found in habitats as
diverse as dry savannahs, 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 ecotonesbetween forests and savannahs. 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, grass and
roots. They are born with sterile intestines and require bacteria obtained
from their mother's feces to digest vegetation. African elephants 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. Elephants average 3–4 hours of sleep per day. 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. Elephants go on seasonal
migrations in search of food, water, minerals, and mates. At Chobe National Park, Botswana, 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 savannah 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; African forest elephants ingest and
defecate seeds, with either no effect or a positive effect on germination. The seeds 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 elephants 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. Their weight can compact the soil, which causes the rain
to run off, leading to erosion.
Elephants do not digest much of their food. Other
animals, such as this baboon, may pick through elephant dung looking for undigested
seeds.
Elephants typically coexist peacefully with other
herbivores, which will usually stay out of 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. The size of adult elephants makes them nearly
invulnerable to predators, though there are rare reports of adult
elephants falling prey to tigers. 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 juvenile elephants
during the dry season, and a pride of 30 lions has been recorded killing
juvenile individuals between the ages of four and eleven years. Elephants
appear to distinguish between the growls of larger predators like tigers and
smaller ones like leopards (which have not been recorded killing calves); the
latter they react less fearfully and more aggressively to. Elephants tend
to have high numbers of parasites, particularly nematodes, compared to other herbivores. This is due to lower
predation pressures that would otherwise kill off many of the individuals with
significant parasite loads.
Social organisation
Female elephants spend their entire lives in
tight-knit matrilineal family groups, some of which are made up of more
than ten members, including three pairs of mothers with offspring, and are led
by the matriarch which is often the eldest female. 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 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, which may be closely
related due to previously being part of the same family group which split after
becoming too large for the available resources. 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.
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 social life of the adult male is very different. As
he matures, a male spends 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. When males
permanently leave, they either live alone or with other males. The former is
typical of bulls in dense forests. Asian males are usually solitary, but
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. These elephants can be quite sociable when not competing for
dominance or mates, and will form long-term relationships. A dominance hierarchy exists among males, whether they range socially or
solitarily. Dominance depends on the age, size and sexual condition, and
when in groups, males follow the lead of the dominant bull. Young bulls may
seek out the company and leadership of older, more experienced
males, whose presence appears to control their aggression and prevent them
from exhibiting "deviant" behaviour. Adult males and females
come together for reproduction. Bulls associate with family groups if an oestrous cow is present.
Sexual behaviour
Musth
Adult males enter a state of increased testosterone known as musth.
In a population in southern India, males first enter musth at the age of 15,
but it is not very intense until they are older than 25. At Amboseli, bulls
under 24 do not go into musth, while half of those aged 25–35 and all those
over 35 do. Young bulls appear to enter musth during the dry season
(January–May), while older bulls go through it during the wet season
(June–December). The main characteristic of a bull's musth is a fluid secreted
from the temporal
gland that runs down the side of his face.
He may urinate with his penis still in his sheath, which causes the urine to spray on his hind legs.
Behaviours associated with musth include walking with the head held high and
swinging, picking at the ground with the tusks, marking, rumbling and waving
only one ear at a time. This can last from a day to four months.
Males become extremely aggressive during musth. Size is
the determining factor in agonistic encounters when the individuals have the same
condition. In contests between musth and non-musth individuals, musth bulls win
the majority of the time, even when the non-musth bull is larger. A male may
stop showing signs of musth when he encounters a musth male of higher rank.
Those of equal rank tend to avoid each other. Agonistic encounters typically
consist of threat displays, chases, and minor sparring with the tusks. Serious
fights are rare.
Mating
Elephants are polygynous breeders, and copulations are most frequent during the peak of the wet
season. A cow in oestrus releases chemical signals (pheromones) in her urine and vaginal secretions to signal her
readiness to mate. A bull will follow a potential mate and assess her condition
with the flehmen
response, which requires the male to collect a
chemical sample with his trunk and bring it to the vomeronasal organ. The oestrous cycle of a cow lasts 14–16 weeks with
a 4–6-week follicular
phase and an 8- to 10-week luteal phase. While most mammals have one surge of luteinizing hormone during the follicular phase, elephants have two.
The first (or anovulatory) surge, could signal to males that the female is in
oestrus by changing her scent, but ovulation does not occur until the second (or ovulatory)
surge. Fertility rates in cows decline around 45–50 years of age.
Bulls engage in a behaviour known as mate-guarding, where
they follow oestrous females and defend them from other males. Most
mate-guarding is done by musth males, and females actively seek to be guarded
by them, particularly older ones. Thus these bulls have more reproductive
success. Musth appears to signal to females the condition of the male, as
weak or injured males do not have normal musths. For young females, the
approach of an older bull can be intimidating, so her relatives stay nearby to
provide support and reassurance. During copulation, the male lays his
trunk over the female's back. The penis is very mobile, being able to move
independently of the pelvis. Prior to mounting, it curves forward and
upward. Copulation lasts about 45 seconds and does not involve pelvic thrusting or ejaculatory pause. Elephant sperm must swim
close to 2 m (6.6 ft) to reach the egg. By comparison, human sperm
only has to swim around 76.2 mm (3.00 in).
Homosexual
behaviour is frequent in both sexes. As in
heterosexual interactions, this involves mounting. Male elephants sometimes
stimulate each other by playfighting and "championships" may form
between old bulls and younger males. Female same-sex behaviours have been
documented only in captivity where they are known to masturbate
one another with their trunks.
Birthing and calves
Gestation in elephants typically lasts around two years 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). Typically, only a single young
is born, but twins sometimes occur. The relatively long pregnancy is
maintained by five 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. 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. Allomothers are typically two
to twelve years old. When a predator is near, the family group gathers
together with the calves in the centre.
For
the first few days, the newborn is unsteady on its feet, 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. It still needs its mother 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
rate, body condition and reproductive ability. Play behaviour in calves
differs between the sexes; females run or chase each other while males
play-fight. The former are sexually mature by the age of nine years while the latter
become mature around 14–15 years. Adulthood starts at about 18 years of age in
both sexes. Elephants have long lifespans, reaching 60–70 years of
age. Lin Wang, a captive male Asian elephant, lived for 86 years.
Communication
Touching
is an important form of communication among elephants. Individuals 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 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 several sounds, usually through
the larynx, though some may be modified by the trunk. Perhaps the most well known call
is the trumpet which is made by blowing through the trunk. Trumpeting is made
during excitement, distress or aggression. Fighting elephants may roar or
squeal, and wounded ones may bellow. Rumbles are produced during mild arousal and some
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
with sound pressure levels as high 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 the matriarch 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 possess 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
individual running or mock charging can create seismic signals that can be
heard at great distances. When detecting the seismics of an alarm call
signalling danger from predators, elephants enter a defensive posture and
family groups will pack together. Seismic waveforms produced by locomotion
appear to travel distances of up to 32 km (20 mi) while those from
vocalisations travel 16 km (10 mi).
Intelligence and cognition
Elephants exhibit mirror self-recognition, an indication of self-awareness and cognition that has also been demonstrated in some apesand dolphins. One study of a captive female Asian elephant
suggested the animal was capable of learning and distinguishing between several
visual and some acoustic discrimination pairs. This individual was even able to
score a high accuracy rating when re-tested with the same visual pairs a year
later Elephants are among the species known to use
tools. An Asian elephant has been observed
modifying branches and using them as flyswatters. Tool modification by these animals is not as
advanced as that of 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. 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 show interest in the bones of their own
kind, regardless of whether they are related. As with chimps and dolphins,
a dying or dead elephant may elicit attention and aid from others, including
those from other groups. This has been interpreted as expressing
"concern"; however, others would dispute such an interpretation
as being anthropomorphic; the Oxford Companion to Animal Behaviour (1987) advised that "one is well advised to
study the behaviour rather than attempting to get at any underlying
emotion".
Conservation
Status
Distribution of elephants
African elephants were listed as vulnerable by the International
Union for Conservation of Nature (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. By
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, fewer than had previously been thought. From 1977 to 1989,
elephant populations declined by 74% in East Africa. After 1987, losses in
elephant numbers accelerated, and savannah populations from Cameroon to Somalia experienced 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%. Due
to the vast areas involved, assessing the total African elephant population
remains difficult and involves an element of guesswork. The IUCN estimates a
total of around 440,000 individuals for 2012.
African elephants receive at least some legal protection
in every country where they are found, but 70% of their range exists outside
protected areas. Successful conservation efforts in certain areas have led to
high population densities. As of 2008, local numbers were controlled by
contraception or translocation. Large-scale cullings ceased in 1988, when Zimbabwe abandoned the
practice. In 1989, the African elephant was listed under Appendix I by
the Convention on
International Trade in Endangered Species of Wild Fauna and Flora (CITES), making trade illegal. Appendix II status
(which allows restricted trade) was given to elephants in Botswana, Namibia,
and Zimbabwe in 1997 and South Africa in 2000. In some countries, sport hunting of the animals is legal; Botswana, Cameroon, Gabon,
Mozambique, Namibia, South Africa, Tanzania, Zambia, and Zimbabwe have CITES export quotas for
elephant trophies. In June 2016, the First Lady of Kenya, Margaret Kenyatta, helped launch the East Africa Grass-Root Elephant
Education Campaign Walk, organised by elephant conservationist Jim Nyamu. The event was conducted to raise awareness of the value
of elephants and rhinos, to help mitigate human-elephant conflicts, and to
promote anti-poaching activities.
In 2008, the IUCN listed the Asian elephant as endangered due to a 50% population decline over the past 60–75
years while CITES lists the species under Appendix I. Asian elephants
once ranged from Syria and Iraq (the
subspecies Elephas maximus asurus), to China (up to the Yellow River) and Java. It is now extinct in these
areas, and the current range of Asian elephants is highly
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 is in India. Although Asian elephants are
declining in numbers overall, particularly in Southeast Asia, the population in
the Western
Ghatsappears to be increasing.
Threats
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
rivalled that of gold. The ivory trade 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 elephant
populations were healthy, but only if their supplies were from elephants that
had been culled or died of natural causes.
The ban allowed the elephant to recover in parts of
Africa. In January 2012, 650 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 in India. China was the biggest
market for poached ivory but announced they would phase out the legal domestic
manufacture and sale of ivory products in May 2015, and in September 2015,
China and the United States said "they would enact a nearly complete ban
on the import and export of ivory" due to causes of extinction.
Other threats to elephants include habitat destruction and fragmentation. 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. 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. One
proposed solution is the provision of ‘urban corridors’ which allow the animals
access to key areas.
Elephants
and humans
Working animal
Elephants have been working animals since at least the Indus Valley
Civilization and continue to be used in
modern times. There were 13,000–16,500 working elephants employed in Asia in
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.
Individuals
of the Asian species have been often trained as working animals. In the forests
of Myanmar, for example, elephants have played a critical role in sustainable
logging. Under the guidance of their handlers, elephants are used to manipulate
and drag heavy teak logs from road-less areas to rivers and roads. From there,
the valuable logs are shipped to commercial mills.
The practice of working elephants has also been attempted
in Africa. The taming of African elephants in the Belgian Congo began by decree of Leopold II of
Belgium during the 19th century and
continues to the present with the Api
Elephant Domestication Centre.
Asian elephants perform tasks such as hauling loads into
remote areas, moving logs to rivers and roads, transporting tourists
around national
parks, pulling wagons, and leading religious
processions. In northern Thailand, the animals are used to digest coffee beans for Black Ivory coffee. They are valued 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. Elephants can be trained to respond to over 30 commands. Musth
bulls can be difficult and dangerous to work with and are chained and
semi-starved until the condition passes. In India, many working elephants
are alleged to have been subject to abuse. They and other captive elephants are
thus protected under The
Prevention of Cruelty to Animals Act of 1960.
In both Myanmar and Thailand, deforestation and other economic factors have resulted in sizable
populations of unemployed elephants resulting in health problems for the
elephants themselves as well as economic and safety problems for the people
amongst whom they live.
Warfare
Historically,
elephants were considered formidable instruments of war. 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 earliest references to war elephants is in the
Indian epic Mahabharata (written
in the 4th century BC, but said to describe events between the 11th and 8th
centuries BC). 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 BC), 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 BC) army while 9,000 may
have been used in the Mauryan army (between the 4th and 2nd centuries BC). The Arthashastra (written around 300 BC) advised the Mauryan
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 BC) while
Southeast Asian states first used war elephants possibly as early as the 5th
century BC 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 BC).
His son and successor Ptolemy II (who began his rule in 285 BC) obtained his supply
of elephants further south in Nubia.
From then on, war elephants were employed in the Mediterranean and North Africa
throughout the classical period. The Greek king Pyrrhus used elephants in his attempted invasion of Rome in
280 BC. While they frightened 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 BC 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 them 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 trained to perform
tricks. The most famous circus elephant was probably Jumbo (1861
– 15 September 1885), who was a major attraction in the Barnum
& Bailey Circus. 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. After
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. As of 2000,
around 1,200 Asian and 700 African elephants were 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.
Keeping elephants in zoos has 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.
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, 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. In January 2016,
Ringling Bros. and Barnum and Bailey circus announced it would retire its
touring elephants in May 2016.
Disease transmission
Like many mammals, elephants can contract and transmit
diseases to humans, one of which is tuberculosis. As of 2015, tuberculosis appears to be widespread among
captive elephants in the US, and because the disease can spread through the air
to infect both humans and other animals, it is a public health concern
affecting circuses and zoos. In
2012, two elephants in Tete d'Or
zoo, Lyon were diagnosed with the disease. Due
to the threat of transmitting tuberculosis to other animals or visitors to the
zoo, their euthanasia was initially ordered by city authorities but a
court later overturned this decision. At an elephant sanctuary in
Tennessee, a 54-year-old African elephant was considered to be the source of
tuberculosis infections among eight workers.
Attacks
Elephants can exhibit bouts of aggressive behaviour and
engage in destructive actions against humans. In Africa, groups of
adolescent elephants damaged homes in villages after cullings in the 1970s and
1980s. Because of the timing, these attacks have been interpreted as
vindictive. In parts of 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.
Cultural depictions
In many cultures, elephants represent strength, power,
wisdom, longevity, stamina, leadership, sociability, nurturance and loyalty.
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.
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 Asia, 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 the 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.[206] 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 of central Africa 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, near the Niger Delta, 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 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
behaviour. They are typically surrogates for humans with 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.