HOME

TheInfoList



OR:

Elephants are the
largest Large means of great size. Large may also refer to: Mathematics * Arbitrarily large, a phrase in mathematics * Large cardinal, a property of certain transfinite numbers * Large category, a category with a proper class of objects and morphisms (o ...
existing land animals. Three living
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are currently recognised: the
African bush elephant The African bush elephant (''Loxodonta africana'') is one of two extant African elephant species and one of three extant elephant species. It is the largest living terrestrial animal, with bulls reaching a shoulder height of up to and a body ...
, the
African forest elephant The African forest elephant (''Loxodonta cyclotis'') is one of the two living African elephant species. It is native to humid forests in West Africa and the Congo Basin. It is the smallest of the three living elephant species, reaching a should ...
, and the
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living species of the genus ''Elephas'' and is distributed throughout the Indian subcontinent and Southeast Asia, from India in the west, Nepal in the no ...
. They are the only surviving members of the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Elephantidae Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are terrestrial large mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the ...
and the order Proboscidea. The order was formerly much more diverse during the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
, but most species became extinct during the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
epoch. Distinctive features of elephants include a long
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
called a trunk,
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s, large ear flaps, pillar-like legs, and tough but sensitive skin. The trunk is used for breathing, bringing food and water to the mouth, and grasping objects. Tusks, which are derived from the incisor teeth, serve both as weapons and as tools for moving objects and digging. The large ear flaps assist in maintaining a constant body temperature as well as in communication. African elephants have larger ears and concave backs, whereas Asian elephants have smaller ears, and convex or level backs. Elephants are scattered throughout
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
,
South Asia South Asia is the southern subregion of Asia, which is defined in both geographical and ethno-cultural terms. The region consists of the countries of Afghanistan, Bangladesh, Bhutan, India, Maldives, Nepal, Pakistan, and Sri Lanka.;;;;;;;; ...
, and
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
and are found in different habitats, including
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
hs, forests, deserts, and
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...
es. They are
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
, and they stay near water when it is accessible. They are considered to be
keystone species A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintaini ...
, due to their impact on their environments. Elephants have a
fission–fusion society In ethology, a fission–fusion society is one in which the size and composition of the social group change as time passes and animals move throughout the environment; animals merge into a group (fusion)—e.g. sleeping in one place—or split (fis ...
, in which multiple family groups come together to socialise. 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, which do not include bulls, are usually led by the oldest cow, known as the
matriarch Matriarchy is a social system in which women hold the primary power positions in roles of authority. In a broader sense it can also extend to moral authority, social privilege and control of property. While those definitions apply in general E ...
. 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. They enter a state of increased
testosterone Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testes and prostate, as well as promoting secondar ...
and aggression known as
musth Musth or must (from Persian, )''The Oxford Dictionary and Thesaurus: American edition'', published 1996 by Oxford University Press; p. 984 is a periodic condition in bull (male) elephants characterized by aggressive behavior and accompanied by ...
, which helps them gain dominance over other males as well as 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 Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inquir ...
by touch, sight, smell, and sound; elephants use
infrasound Infrasound, sometimes referred to as low status sound, describes sound waves with a frequency below the lower limit of human audibility (generally 20 Hz). Hearing becomes gradually less sensitive as frequency decreases, so for humans to perce ...
and
seismic communication Seismic or vibrational communication is a process of conveying information through mechanical (seismic) vibrations of the substrate. The substrate may be the earth, a plant stem or leaf, the surface of a body of water, a spider's web, a honeycomb, ...
over long distances. Elephant intelligence has been compared with that of
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s and
cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
ns. They appear to have
self-awareness In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifesty ...
, and appear to show
empathy Empathy is the capacity to understand or feel what another person is experiencing from within their frame of reference, that is, the capacity to place oneself in another's position. Definitions of empathy encompass a broad range of social, co ...
for dying and dead family members. African bush elephants and Asian elephants are listed as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
and African forest elephants as Critically Endangered by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN). One of the biggest threats to elephant populations is the
ivory trade The ivory trade is the commercial, often illegal trade in the ivory tusks of the hippopotamus, walrus, narwhal, mammoth, and most commonly, African and Asian elephants. Ivory has been traded for hundreds of years by people in Africa and Asia, ...
, as the animals are poached for their ivory tusks. Other threats to wild elephants include
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
and conflicts with local people. Elephants are used as
working animal A working animal is an animal, usually domesticated, that is kept by humans and trained to perform tasks instead of being slaughtered to harvest animal products. Some are used for their physical strength (e.g. oxen and draft horses) or for t ...
s 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
circus A circus is a company of performers who put on diverse entertainment shows that may include clowns, acrobats, trained animals, trapeze acts, musicians, dancers, hoopers, tightrope walkers, jugglers, magicians, ventriloquists, and unicyclist ...
es. Elephants are highly recognisable and have been featured in art, folklore, religion, literature, and
popular culture Popular culture (also called mass culture or pop culture) is generally recognized by members of a society as a set of practices, beliefs, artistic output (also known as, popular art or mass art) and objects that are dominant or prevalent in a ...
.


Etymology

The word "elephant" is based on the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''elephas'' (
genitive In grammar, the genitive case (abbreviated ) is the grammatical case that marks a word, usually a noun, as modifying another word, also usually a noun—thus indicating an attributive relationship of one noun to the other noun. A genitive can al ...
''elephantis'') ("elephant"), which is the Latinised form of the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
ἐλέφας (''elephas'') (genitive ἐλέφαντος (''elephantos''), probably from a non-
Indo-European The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, Dutc ...
language, likely Phoenician. It is attested in
Mycenaean Greek Mycenaean Greek is the most ancient attested form of the Greek language, on the Greek mainland and Crete in Mycenaean Greece (16th to 12th centuries BC), before the hypothesised Dorian invasion, often cited as the ''terminus ad quem'' for the ...
as ''e-re-pa'' (genitive ''e-re-pa-to'') in
Linear B Linear B was a syllabic script used for writing in Mycenaean Greek, the earliest attested form of Greek. The script predates the Greek alphabet by several centuries. The oldest Mycenaean writing dates to about 1400 BC. It is descended from ...
syllabic script. As in Mycenaean Greek,
Homer Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
used the Greek word to mean
ivory Ivory is a hard, white material from the tusks (traditionally from elephants) and teeth of animals, that consists mainly of dentine, one of the physical structures of teeth and tusks. The chemical structure of the teeth and tusks of mammals is ...
, but after the time of
Herodotus Herodotus ( ; grc, , }; BC) was an ancient Greek historian and geographer from the Greek city of Halicarnassus, part of the Persian Empire (now Bodrum, Turkey) and a later citizen of Thurii in modern Calabria ( Italy). He is known f ...
, it also referred to the animal. The word "elephant" appears in
Middle English Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English p ...
as ''olyfaunt'' (c.1300) and was borrowed from
Old French Old French (, , ; Modern French: ) was the language spoken in most of the northern half of France from approximately the 8th to the 14th centuries. Rather than a unified language, Old French was a linkage of Romance dialects, mutually intelligib ...
''oliphant'' (12th century).


Taxonomy

Alongside their closest relatives belong to the family
Elephantidae Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are terrestrial large mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the ...
, the sole remaining family within the order Proboscidea. Their closest
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
relatives are the
sirenia The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct ...
ns (
dugong The dugong (; ''Dugong dugon'') is a marine mammal. It is one of four living species of the order Sirenia, which also includes three species of manatees. It is the only living representative of the once-diverse family Dugongidae; its closest m ...
s and
manatee Manatees (family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing three of the four living species ...
s) and the
hyrax Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...
es, with which they share the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Paenungulata Paenungulata (from Latin ''paene'' "almost" + ''ungulātus'' "having hoofs") is a clade of "sub-ungulates", which groups three extant mammal orders: Proboscidea (including elephants), Sirenia (sea cows, including dugongs and manatees), and H ...
within the superorder
Afrotheria Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a clade of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews (also k ...
. Elephants and sirenians are further grouped in the clade
Tethytheria Tethytheria is a clade of mammals that includes the sirenians and proboscideans, as well as the extinct order Embrithopoda. Though there is strong anatomical and molecular support for the monophyly of Tethytheria, the interrelationships between ...
. Three species of elephants are recognised; the
African bush elephant The African bush elephant (''Loxodonta africana'') is one of two extant African elephant species and one of three extant elephant species. It is the largest living terrestrial animal, with bulls reaching a shoulder height of up to and a body ...
('' Loxodonta africana'') and
forest elephant The African forest elephant (''Loxodonta cyclotis'') is one of the two living African elephant species. It is native to humid forests in West Africa and the Congo Basin. It is the smallest of the three living elephant species, reaching a shoulde ...
(''Loxodonta cyclotis'') of
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
, and the
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living species of the genus ''Elephas'' and is distributed throughout the Indian subcontinent and Southeast Asia, from India in the west, Nepal in the no ...
(''
Elephas ''Elephas'' is one of two surviving genera in the family of elephants, Elephantidae, with one surviving species, the Asian elephant, ''Elephas maximus''. Several extinct species have been identified as belonging to the genus, extending back to ...
maximus'') of
South South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
and
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
. 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 The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
are narrower in the Asian elephant while those of the African are more diamond-shaped. The Asian elephant also has
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
bumps on its head and some patches of
depigmentation Depigmentation is the lightening of the skin or loss of pigment. Depigmentation of the skin can be caused by a number of local and systemic conditions. The pigment loss can be partial (injury to the skin) or complete (caused by vitiligo). It can be ...
on its skin.Shoshani, pp. 38–41. Among African elephants, forest elephants have smaller and more rounded ears and thinner and straighter tusks than bush elephants and are limited in range to the forested areas of
western Western may refer to: Places *Western, Nebraska, a village in the US *Western, New York, a town in the US *Western Creek, Tasmania, a locality in Australia *Western Junction, Tasmania, a locality in Australia *Western world, countries that id ...
and
Central Africa Central Africa is a subregion of the African continent comprising various countries according to different definitions. Angola, Burundi, the Central African Republic, Chad, the Democratic Republic of the Congo, the Republic of the Congo, ...
. Both were traditionally considered a single species, ''Loxodonta africana'', but molecular studies have affirmed their status as separate species. In 2017, DNA sequence analysis showed that ''L. cyclotis'' is more closely related to the extinct ''
Palaeoloxodon antiquus The straight-tusked elephant (''Palaeoloxodon antiquus'') is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene (781,000–30,000 years before present). Recovered individuals have reac ...
'', than it is to ''L. africana,'' possibly undermining the genus ''Loxodonta'' as a whole.


Evolution and extinct relatives

Over 180 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 The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
, heralded the first radiation. The
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
included ''
Numidotherium ''Numidotherium'' ("Numidia beast") is an extinct genus of early proboscideans, discovered in 1984, that lived during the middle Eocene of North Africa some 46 million years ago. It was about 90-100 cm tall at the shoulder and weighed about 250- ...
'', ''
Moeritherium ''Moeritherium'' ("the beast from Lake Moeris") is an extinct genus of primitive proboscideans. These prehistoric mammals are related to the elephant and, more distantly, sea cows and hyraxes. They lived during the Eocene epoch. Description ''M ...
,'' and ''
Barytherium ''Barytherium'' (meaning "heavy beast") is a genus of an extinct family (Barytheriidae) of primitive proboscideans that lived during the late Eocene and early Oligocene in North Africa. The type species is ''Barytherium grave'', found at the ...
'' from Africa. These animals were relatively small and aquatic. Later on, genera such as ''
Phiomia ''Phiomia'' is an extinct genus of basal proboscid that lived in what is now Northern Africa during the Late Eocene to Early Oligocene some 37–30 million years ago.Strauss, BPrehistoric Elephant Pictures and Profiles/ref> "''Phiomia serridens''" ...
'' and ''
Palaeomastodon ''Palaeomastodon'' an extinct genus of Proboscidea. ''Palaeomastodon'' fossils have been found in Africa, where they lived some 36-35 million years ago. They are believed to be the ancestors of elephants or mastodons. ''Palaeomastodon'' lived in ...
'' arose; the latter likely inhabited forests and open woodlands. Proboscidean diversity declined during the Oligocene. One notable species of this epoch was ''
Eritreum ''Eritreum melakeghebrekristosi'' is an extinct species of proboscidean mammal, which lived in Northeast Africa during the late Oligocene some 27 million years ago, and is considered to be the missing link between modern elephants and their ance ...
melakeghebrekristosi'' of the
Horn of Africa The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004), ...
, which may have been an ancestor to several later species. The beginning of the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
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,Sukumar, pp. 16–19. 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 ''Gomphotherium'' (; "welded beast") is an extinct genus of proboscids from the Neogene and early Pleistocene of Eurasia, Africa, North America and Asia. As of 2021, two species, ''G. annectens'' and possibly ''G. subtapiroideum'', are also k ...
'' and ''
Platybelodon ''Platybelodon'' ("flat-spear tusk") is an extinct genus of large herbivorous proboscidean mammals related to modern-day elephants. Species lived during the middle Miocene Epoch in Africa, Asia and the Caucasus. Palaeobiology ''Platybelodon'' w ...
''. 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 ''Primelephas'' is a genus of Elephantinae that existed during the Miocene and Pliocene epochs. The name of the genus suggests 'first elephant'. These primitive elephantids are thought to be the common ancestor of ''Mammuthus'', the mammoths ...
gomphotheroides'' gave rise to ''Loxodonta'', ''Mammuthus,'' and ''Elephas''. ''Loxodonta'' branched off earliest around the Miocene and
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58stegodontids, 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 American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, p ...
. At the beginning of the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
, elephantids experienced a high rate of
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. The Pleistocene also saw the arrival of ''
Palaeoloxodon namadicus ''Palaeoloxodon namadicus'' or the Asian straight-tusked elephant, is an extinct species of prehistoric elephant known from the early Middle to Late Pleistocene of the Indian subcontinent, and possibly also elsewhere in Asia. Some authorities r ...
'', the largest terrestrial mammal of all time. ''
Loxodonta atlantica ''Loxodonta atlantica'' is an extinct species of elephant in the genus '' Loxodonta'', from Africa. It was larger than the modern African elephant, with more progressive dentition. It includes Pleistocene fossils from Ternifine, Middle Pleisto ...
'' 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.Sukumar, p. 44. ''Mammuthus'' evolved into several species, including the well-known
woolly mammoth The woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived during the Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with '' Mammuthus s ...
.Sukumar, pp. 28–31. Interbreeding appears to have been common among elephantid species, which in some cases led to species with three ancestral genetic components, such as the ''Palaeoloxodon antiquus''. In the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
, most proboscidean species vanished during the
Quaternary glaciation The Quaternary glaciation, also known as the Pleistocene glaciation, is an alternating series of glacial and interglacial periods during the Quaternary period that began 2.58 Ma (million years ago) and is ongoing. Although geologists describe ...
which
killed off The killing off of a character is a device in fiction, whereby a character dies, but the story continues. The term, frequently applied to television, film, video game, anime, manga and chronological series, often denotes an untimely or unexpect ...
50% of genera weighing over worldwide. Proboscideans experienced several evolutionary trends, such as an increase in size, which led to many giant species that stood up to tall. As with other
megaherbivore Megaherbivores (Greek μέγας megas "large" and Latin ''herbivora'' "herbivore") are large terrestrial herbivores that can exceed in weight. This polyphyletic group of megafauna includes elephants, rhinos, hippos, and giraffes. The largest ...
s, including the extinct
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s, 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 In anatomy, a sesamoid bone () is a bone embedded within a tendon or a muscle. Its name is derived from the Arabic word for ' sesame seed', indicating the small size of most sesamoids. Often, these bones form in response to strain, or can be pres ...
providing support. Early proboscideans developed longer
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s and smaller craniums while more
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
ones developed shorter mandibles, which shifted the head's
centre of gravity In physics, the center of mass of a distribution of mass in space (sometimes referred to as the balance point) is the unique point where the weighted relative position of the distributed mass sums to zero. This is the point to which a force may ...
. 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
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s, incisors and
canines Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
decreased. The cheek teeth (molars and premolars) of proboscideans 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 The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
anatomy. Several species of proboscideans lived on islands and experienced
insular dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
. This occurred primarily during the Pleistocene when some elephant populations became isolated by fluctuating sea levels, although
dwarf elephant Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes (around ) in comparison with their immediate ancestors. Dwarf elephants are an example ...
s 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
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
s develop
gigantism Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...
in these conditions. Dwarf elephants are known to have lived in
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
, the
Channel Islands of California The Channel Islands () are an eight-island archipelago located within the Southern California Bight in the Pacific Ocean, off the coast of California. The four Northern Channel Islands are part of the Transverse Ranges geologic province, and ...
, and several islands of the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
.Sukumar, pp. 31–33.


Anatomy


Size

Elephants are the largest living terrestrial animals. African bush elephants are the largest species, with males being tall at the shoulder with a body mass of and females standing tall at the shoulder with a body mass of . Male Asian elephants are usually about tall at the shoulder and whereas females are tall at the shoulder and . African forest elephants are the smallest species, with males usually being around tall at the shoulder and . Male African bush elephants are typically 23% taller than females, whereas male Asian elephants are only around 15% taller than females.


Bones

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.


Head

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 A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
in every direction. The skull contains air cavities (
sinuses Paranasal sinuses are a group of four paired air-filled spaces that surround the nasal cavity. The maxillary sinuses are located under the eyes; the frontal sinuses are above the eyes; the ethmoidal sinuses are between the eyes and the sphenoid ...
) that reduce the weight of the skull while maintaining overall strength. These cavities give the inside of the skull a
honeycomb A honeycomb is a mass of Triangular prismatic honeycomb#Hexagonal prismatic honeycomb, hexagonal prismatic Beeswax, wax cells built by honey bees in their beehive, nests to contain their larvae and stores of honey and pollen. beekeeping, Beekee ...
-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.Shoshani, pp. 68–70. Because of the size of the head, the neck is relatively short to provide better support. Lacking a
lacrimal apparatus The lacrimal apparatus is the physiological system containing the Orbit (anatomy), orbital structures for tears, tear production and drainage.Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Com ...
, the eye relies on the
harderian gland The Harderian gland is a gland found within the eye's orbit that occurs in tetrapods (reptiles, amphibians, birds and mammals) that possess a nictitating membrane. The gland can be compound tubular or compound tubuloalveolar, and the fluid it s ...
to keep it moist. A durable
nictitating membrane The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
protects the eye globe. The animal's
field of vision The visual field is the "spatial array of visual sensations available to observation in introspectionist psychological experiments". Or simply, visual field can be defined as the entire area that can be seen when an eye is fixed straight at a point ...
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.


Ears

Elephant ears have thick bases with thin tips. The ear flaps, or
pinnae The auricle or auricula is the visible part of the ear that is outside the head. It is also called the pinna (Latin for "wing" or " fin", plural pinnae), a term that is used more in zoology. Structure The diagram shows the shape and location ...
, contain numerous blood vessels called
capillaries A capillary is a small blood vessel from 5 to 10 micrometres (μm) in diameter. Capillaries are composed of only the tunica intima, consisting of a thin wall of simple squamous endothelial cells. They are the smallest blood vessels in the body: ...
. 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 The hertz (symbol: Hz) is the unit of frequency in the International System of Units (SI), equivalent to one event (or cycle) per second. The hertz is an SI derived unit whose expression in terms of SI base units is s−1, meaning that on ...
(in close proximity to the
Soprano A soprano () is a type of classical female singing voice and has the highest vocal range of all voice types. The soprano's vocal range (using scientific pitch notation) is from approximately middle C (C4) = 261  Hz to "high A" (A5) = 880&n ...
C).


Trunk

The trunk, or
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
, is a fusion of the nose and upper lip, although in early
fetal A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal develo ...
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 fascicle A muscle fascicle is a bundle of skeletal muscle fibers surrounded by perimysium, a type of connective tissue. Structure Muscle cells are grouped into muscle fascicles by enveloping perimysium connective tissue. Fascicles are bundled together b ...
s, 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 Transverse may refer to: *Transverse engine, an engine in which the crankshaft is oriented side-to-side relative to the wheels of the vehicle *Transverse flute, a flute that is held horizontally * Transverse force (or ''Euler force''), the tangen ...
and
radiating In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visib ...
muscles. The muscles of the trunk connect to a bony opening in the skull. The
nasal septum The nasal septum () separates the left and right airways of the Human nose, nasal cavity, dividing the two nostrils. It is Depression (kinesiology), depressed by the depressor septi nasi muscle. Structure The fleshy external end of the nasal ...
is composed of tiny muscle units that stretch horizontally between the nostrils.
Cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck an ...
divides the nostrils at the base.Shoshani, pp. 74–77. As a
muscular hydrostat A muscular hydrostat is a biological structure found in animals. It is used to manipulate items (including food) or to move its host about and consists mainly of muscles with no skeletal support. It performs its hydraulic movement without fluid in ...
, 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 nerve The facial nerve, also known as the seventh cranial nerve, cranial nerve VII, or simply CN VII, is a cranial nerve that emerges from the pons of the brainstem, controls the muscles of facial expression, and functions in the conveyance of tas ...
s – runs along both sides of the trunk. Elephant trunks have multiple functions, including breathing,
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
, touching, grasping, and sound production. The animal's sense of smell may be four times as sensitive as that of a
bloodhound The bloodhound is a large scent hound, originally bred for hunting deer, wild boar and, since the Middle Ages, for tracking people. Believed to be descended from hounds once kept at the Abbey of Saint-Hubert, Belgium, in French it is called, ''l ...
. The trunk's ability to make powerful twisting and coiling movements allows it to collect food, wrestle with other elephants,Kingdon, p. 9. and lift up to . 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 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 are capable of dilating their nostrils at a radius of nearly 30%, increasing the nasal volume by 64%, and can inhale at over which is around 30 times the speed of a human sneeze. Elephants can suck up food and water both to spray in the mouth and, in the case of the latter, to sprinkle on their bodies. An adult Asian elephant is capable of holding 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 Floppy trunk syndrome (abbreviated FTS, also known as flaccid trunk paralysis) is a condition that causes trunk paralysis in African bush elephants. Initially observed in 1989, the syndrome primarily affected bull elephants in several select region ...
is a condition of trunk
paralysis Paralysis (also known as plegia) is a loss of motor function in one or more muscles. Paralysis can also be accompanied by a loss of feeling (sensory loss) in the affected area if there is sensory damage. In the United States, roughly 1 in 50 ...
in African bush elephants caused by the degradation of the
peripheral nerves The peripheral nervous system (PNS) is one of two components that make up the nervous system of bilateral animals, with the other part being the central nervous system (CNS). The PNS consists of nerves and ganglia, which lie outside the brain an ...
and muscles beginning at the tip.


Teeth

Elephants usually have 26 teeth: the
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s, known as the
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s, 12
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s, and 12
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
. Unlike most mammals, which grow baby teeth and then replace them with a single permanent set of adult teeth, elephants are
polyphyodont A polyphyodont is any animal whose teeth are continually replaced. In contrast, diphyodonts are characterized by having only two successive sets of teeth. Polyphyodonts include most toothed fishes, many reptiles such as crocodiles and geckos, and ...
s 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 at four to six years old. The third set falls out at 9–15 years of age and set four lasts until 18–28 years of age. The fifth set of teeth falls out at the 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.Shoshani, pp. 70–71.


Tusks

The tusks of an elephant are modified second incisors in the upper jaw. They replace deciduous
milk teeth Deciduous teeth or primary teeth, also informally known as baby teeth, milk teeth, or temporary teeth,Illustrated Dental Embryology, Histology, and Anatomy, Bath-Balogh and Fehrenbach, Elsevier, 2011, page 255 are the first set of teeth in the ...
at 6–12 months of age and grow continuously at about a year. A newly developed tusk has a smooth enamel cap that eventually wears off. The
dentin Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
e is known as
ivory Ivory is a hard, white material from the tusks (traditionally from elephants) and teeth of animals, that consists mainly of dentine, one of the physical structures of teeth and tusks. The chemical structure of the teeth and tusks of mammals is ...
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 Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
. Much of the tusk can be seen outside; the rest is in a socket in the skull. At least one-third of the tusk contains the
pulp Pulp may refer to: * Pulp (fruit), the inner flesh of fruit Engineering * Dissolving pulp, highly purified cellulose used in fibre and film manufacture * Pulp (paper), the fibrous material used to make paper * Molded pulp, a packaging material ...
and 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 , 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 . In the Asian species, only the males have large tusks. Female Asians have very small tusks, or none at all.Shoshani, pp. 71–74. 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 long and weighed . Hunting for elephant ivory in Africa and Asia has led to
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charle ...
for shorter tusks and tusklessness.


Skin

An elephant's skin is generally very tough, at thick on the back and parts of the head. The skin around the mouth,
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
, 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 A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
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.Shoshani, pp. 66–67. Their hair is thought to be for
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
, helping them lose heat in their hot environments. An elephant uses mud as a sunscreen, protecting its skin from
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
light. Although tough, an elephant's skin is very sensitive. Without regular
mud bath A mud bath is a bath of mud, commonly found in areas where hot spring water can combine with volcanic ash. Mud baths have a long history that dates back thousands of years. Mud baths are conceived as public bathing spaces created in open areas ...
s 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 The surface-area-to-volume ratio, also called the surface-to-volume ratio and variously denoted sa/vol or SA:V, is the amount of surface area per unit volume of an object or collection of objects. SA:V is an important concept in science and engin ...
, 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 A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, and ...
in place of medullary cavities. This strengthens the bones while still allowing
haematopoiesis Haematopoiesis (, from Greek , 'blood' and 'to make'; also hematopoiesis in American English; sometimes also h(a)emopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cell ...
.Shoshani, pp. 69–70. 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 The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
and
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
are fixed in
pronation Motion, the process of movement, is described using specific anatomical terms. Motion includes movement of organs, joints, limbs, and specific sections of the body. The terminology used describes this motion according to its direction relative ...
; the "palm" of the manus faces backward. The pronator quadratus and 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 The giant panda (''Ailuropoda melanoleuca''), also known as the panda bear (or simply the panda), is a bear species endemic to China. It is characterised by its bold black-and-white coat and rotund body. The name "giant panda" is sometimes us ...
'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 The trot is a ten-beat diagonal horse gait where the diagonal pairs of legs move forward at the same time with a moment of suspension between each beat. It has a wide variation in possible speeds, but averages about . A very slow trot is someti ...
,
jump Jumping is a form of locomotion or movement in which an organism or non-living (e.g., robotic) mechanical system propels itself through the air along a ballistic trajectory. Jump or Jumping also may refer to: Places * Jump, Kentucky or Jump S ...
, or
gallop The canter and gallop are variations on the fastest gait that can be performed by a horse or other equine. The canter is a controlled three-beat gait, while the gallop is a faster, four-beat variation of the same gait. It is a natural gait pos ...
. 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 . At this speed, most other
quadrupeds Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
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 swum as far as at a stretch and at speeds of up to .


Organs

The brain of an elephant weighs compared to for a human brain. It is the largest of all terrestrial mammals. While the elephant brain is larger overall, it is proportionally smaller than the human brain when compared to their size. At birth, an elephant's brain already weighs 30–40% of its adult weight. The
cerebrum The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. In ...
and
cerebellum The cerebellum (Latin for "little brain") is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebel ...
are well developed, and the
temporal lobe The temporal lobe is one of the four Lobes of the brain, major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain. The temporal lobe ...
s are so large that they bulge out laterally. Their temporal lobes are proportionally larger than in other animals, including humans. The throat of an elephant appears to contain a pouch where it can store water for later use. The
larynx The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
of the elephant is the largest known among mammals. The
vocal fold The human voice consists of sound made by a human being Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and exceptional cognitive skills due to a large and compl ...
s are long and are attached close to the
epiglottis The epiglottis is a leaf-shaped flap in the throat that prevents food and water from entering the trachea and the lungs. It stays open during breathing, allowing air into the larynx. During swallowing, it closes to prevent aspiration of food int ...
base. When comparing an elephant's vocal folds to those of a human, an elephant's are longer, thicker, and have a larger cross-sectional area. In addition, they are tilted at 45 degrees and positioned more anteriorly than a human's vocal folds. The heart of an elephant weighs . It has a double-pointed
apex The apex is the highest point of something. The word may also refer to: Arts and media Fictional entities * Apex (comics), a teenaged super villainess in the Marvel Universe * Ape-X, a super-intelligent ape in the Squadron Supreme universe *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.Shoshani, pp. 78–79.
Connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
exists in place of the
pleural cavity The pleural cavity, pleural space, or interpleural space is the potential space between the pleurae of the pleural sac that surrounds each lung. A small amount of serous pleural fluid is maintained in the pleural cavity to enable lubrication bet ...
. 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 Hindgut fermentation is a digestive process seen in monogastric herbivores, animals with a simple, single-chambered stomach. Cellulose is digested with the aid of symbiotic bacteria.penis A penis (plural ''penises'' or ''penes'' () is the primary sexual organ that male animals use to inseminate females (or hermaphrodites) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males do n ...
can reach a length of and a diameter of at the base. It is S-shaped when fully erect and has a Y-shaped
orifice An orifice is any opening, mouth, hole or vent, as in a pipe, a plate, or a body * Body orifice, any opening in the body of a human or animal *Orifice plate, a restriction used to measure flow or to control pressure or flow, sometimes given specia ...
. The female has a well-developed
clitoris The clitoris ( or ) is a female sex organ present in mammals, ostriches and a limited number of other animals. In humans, the visible portion – the glans – is at the front junction of the labia minora (inner lips), above the ope ...
at up to . 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 Temporin, 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 Musth or must (from Persian, )''The Oxford Dictionary and Thesaurus: American edition'', published 1996 by Oxford University Press; p. 984 is a periodic condition in bull (male) elephants characterized by aggressive behavior and accompanied by ...
. Females have also been observed with secretions from the temporal glands.Sukumar, p. 149.


Body temperature

Elephants are homeothermy, homeotherms, and maintain their average body temperature at ~ 36 °C, with minimum 35.2 °C during cool season, and maximum 38.0 °C during hot dry season. Sweat glands are absent in the elephant's skin, but water diffuses through the skin, allowing cooling by evaporative loss. Other physiological or behavioral features may assist with thermoregulation such as flapping ears, mud bathing, spraying water on the skin, seeking shade, and adopting different walking patterns. In addition, the interconnected crevices in the elephant's skin is thought to impede dehydration and improve thermal regulation over a long period of time.


Behaviour and life history


Ecology and activities

The African bush elephant can be found in habitats as diverse as dry
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
hs, deserts,
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...
es, and lake shores, and in elevations from sea level to mountains above the snow line. Forest elephants mainly live in Atlantic Equatorial coastal forests, equatorial forests but will enter gallery forests and ecotones between forests and savannahs.Shoshani, pp. 42–51. Asian elephants prefer areas with a mix of grasses, low woody plants, and trees, primarily inhabiting dry Deccan thorn scrub forests, thorn-scrub forests in southern India and Sri Lanka and evergreen forests in Malay Peninsula, Malaya. Elephants are Herbivore, 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 Browsing (herbivory), browsers while Asian elephants are mainly Grazing, grazers. They can consume as much as of food and 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 a day, but distances as far as have been recorded in the Etosha National Park, Etosha region of Namibia. Elephants go on seasonal migrations in search of food, water, minerals, and mates. At Chobe National Park, Botswana, herds travel 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 A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintaini ...
. 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 dispersal, 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 Malayan tapir, tapir. Because most of the food elephants eat goes undigested, their dung can provide food for other animals, such as dung beetles and monkeys.Shoshani, pp. 226–29. 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 surface runoff, run off, leading to erosion.Eltringham, pp. 124–27. 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–iMfolozi Park, 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. Calves may be preyed on by lions, spotted hyenas, and Lycaon pictus, wild dogs in Africa and tigers in Asia. The lions of Chobe National Park#Geography and ecosystems, Savuti, Botswana, have adapted to hunting elephants, mostly calves, juveniles or even sub-adults, during the dry season, and a pride of 30 lions has been normally recorded killing juvenile individuals between the ages of four and eleven years, and a young bull of about 15 years in an exceptional case. There are rare reports of adult Asian elephants falling prey to tigers. Elephants appear to distinguish between the growls of larger predators like tigers and smaller predators like leopards (which have not been recorded killing calves); they react to leopards less fearfully and more aggressively. 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 Matrilineality, matrilineal family groups, some of which are made up of more than ten members, including three mothers and their dependent offspring, and are led by the
matriarch Matriarchy is a social system in which women hold the primary power positions in roles of authority. In a broader sense it can also extend to moral authority, social privilege and control of property. While those definitions apply in general E ...
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;Kingdon, p. 53. 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. One study found that younger matriarchs are more likely than older ones to under-react to severe danger. Family groups may split after becoming too large for the available resources. 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 which 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.Sukumar, pp. 175–79. 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. Bulls only return to the herd to breed or to socialize, they do not provide prenatal care to their offspring but rather play a fatherly role to younger bulls to show dominance. Male 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 Estrous cycle, oestrous cow is present.Sukumar, pp. 179–83.


Sexual behaviour


Musth

Adult males enter a state of increased
testosterone Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testes and prostate, as well as promoting secondar ...
known as
musth Musth or must (from Persian, )''The Oxford Dictionary and Thesaurus: American edition'', published 1996 by Oxford University Press; p. 984 is a periodic condition in bull (male) elephants characterized by aggressive behavior and accompanied by ...
. 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 Temporin, temporal gland that runs down the side of his face. He may urinate with his penis still in his penile sheath, 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 behaviour, 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.Sukumar, pp. 100–08.


Mating

Elephants are Polygyny in animals, polygynous breeders, and animal sexual behaviour, 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. Before 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 to reach the egg. By comparison, human sperm has to swim around only . Homosexual behavior in animals#Elephants, 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 Animal sexual behaviour#Autoeroticism or masturbation, masturbate one another with their trunks.


Birth and development

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 tall and weigh around .Moss, pp. 106–13. 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. 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.Sukumar, pp. 126–29. 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 vocalisations, usually through the larynx, though some may be Vocal resonation, modified by the trunk.Payne and Langbauer, pp. 120–21. These include trumpets, roars, barks, snorts, growls and rumbles which may be produced for either short or long range communication. Elephants may produce Infrasound, infrasonic rumbles. For Asian elephants, these calls have a frequency of 14–24 Hertz, Hz, with sound pressure levels of 85–90 Decibel, dB and last 10–15 seconds. For African elephants, calls range from 15 to 35 Hz with sound pressure levels as high as 117 dB, allowing communication for many kilometres, with a possible maximum range of around . Elephants are known to seismic communication, communicate with seismics, vibrations produced by impacts on the earth's surface or acoustical waves that travel through it. An individual running or mock charging can create seismic signals that can be heard at travel distances of up to . Seismic waveforms produced from predator alarm calls travel .


Intelligence and cognition

Elephants exhibit Mirror test, mirror self-recognition, an indication of
self-awareness In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifesty ...
and cognition that has also been demonstrated in some apes and 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 Tool use by animals, species known to use tools. An Asian elephant has been observed modifying branches and using them as Fly-killing device, flyswatters. Tool modification by these animals is not as advanced as that of Pan (genus), 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 chimpanzees 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

African bush elephants were listed as Endangered by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN) in 2021, and African forest elephants were listed as Critically Endangered in the same year. 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 East Africa, eastern Africa, 204,000 in southern Africa, and 19,000 in West Africa, 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%. The IUCN estimated that total population in Africa is estimated at around to 415,000 individuals for both species combined as of 2016. 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 Species translocation, translocation. Large-scale cullings ceased in 1988 when Zimbabwe abandoned the practice. In 1989, the African elephant was listed under Appendix I by the CITES, 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, Trophy hunting, 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.Star website
"First Lady launches campaign to save elephants."
4 June 2016. Accessed 5 June 2016.
In 2020, the IUCN listed the Asian elephant as endangered due to an almost 50% population decline over "the last three generations". Asian elephants once ranged from Syria and Iraq (the subspecies Syrian elephant, ''Elephas maximus asurus''), to China (up to the Yellow River)Daniel, p. 174. 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. Around 60% of the population is in India. Although Asian elephants are declining in numbers overall, particularly in Southeast Asia, the population in the Western Ghats appears 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.Martin, pp. 202–07 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 Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
and Habitat fragmentation, fragmentation. The Asian elephant lives in areas with some of the highest human populations and may be confined to small islands of forest among human-dominated landscapes. 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 protection of wildlife corridors which gave the animals greater space and maintain the long term viability of large populations.


Association with humans


Working animal

Elephants have been
working animal A working animal is an animal, usually domesticated, that is kept by humans and trained to perform tasks instead of being slaughtered to harvest animal products. Some are used for their physical strength (e.g. oxen and draft horses) or for t ...
s 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 Mela shikar, traditionally captured with traps and lassos, but since 1950, sedative, tranquillisers have been used. Individuals of the Asian species have been often trained as working animals. 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.McNeely, pp. 149–50. 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,1960, 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. 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 Center, Api Elephant Domestication Centre.


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 Nanda Empire, Nandas (5th and 4th centuries BC) army while 9,000 may have been used in the Maurya Empire, 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 Iran, 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. In his 326 B.C. Indian campaign, Alexander the Great confronted elephants for the first time and suffered heavy casualties. Among the reasons for the refusal of the rank-and-file Macedonian soldiers to continue the Indian conquest were rumors of even larger elephant armies in India. Alexander trained his foot soldiers to injure the animals and cause them to panic during wars with both the Persians and Indians. Ptolemy I Soter, 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 Philadelphus, 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 antiquity, classical period. The Greek king Pyrrhus of Epirus, 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 Carthage, 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.Wylie (2000), pp. 146–48. Overall, elephants owed their initial successes to the element of surprise and to the fear that their great size invoked. With time, strategists devised counter-measures and war elephants turned into an expensive liability and were hardly ever used by Romans and Parthian Empire, Parthians.


Zoos and circuses

Elephants were historically kept for display in the menageries of Ancient Egypt, Elephants in ancient China, China, Ancient Greece, Greece, and Ancient Rome, Rome. The Romans in particular pitted them against humans and other animals in gladiator events. In the Modern history, modern era, elephants have traditionally been a major part of zoos and
circus A circus is a company of performers who put on diverse entertainment shows that may include clowns, acrobats, trained animals, trapeze acts, musicians, dancers, hoopers, tightrope walkers, jugglers, magicians, ventriloquists, and unicyclist ...
es 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 Ringling Bros. and Barnum & Bailey Circus, 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.Tuttle, pp. 184–88. 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 Stereotypy (non-human), 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 Jeffrey Feld, Kenneth Feld acknowledged that circus elephants are struck behind their ears, under their chins and on their legs with metal-tipped prods, called Elephant goad, 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. Ringling Bros. and Barnum and Bailey circus retired its touring elephants in May 2016.


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 can be observed from 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 Motif (visual arts), motifs in Hinduism, Hindu and Buddhism, Buddhist shrines and temples. Elephants were often difficult to portray by people with no first-hand experience of them. The Ancient Rome, 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 architecture, 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 Surrealism, 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.McNeely, pp. 158–60. Similar ideas existed among other African societies, who believed that their chiefs would be reincarnation, 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 Totemism, 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, Gautama Buddha, Buddha is said to have been a White elephant (animal), 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 Popular culture (also called mass culture or pop culture) is generally recognized by members of a society as a set of practices, beliefs, artistic output (also known as, popular art or mass art) and objects that are dominant or prevalent in a ...
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 Republican Party (United States), U.S. Republican Party began with an Republican elephant, 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'', The Walt Disney Company, Disney's ''Dumbo,'' and Kathryn and Byron Jackson's ''The Saggy Baggy Elephant''. Other elephant heroes Anthropomorphism, given human qualities include Jean de Brunhoff's Babar the Elephant, Babar, David McKee's Elmer the Patchwork Elephant, Elmer, and Dr. Seuss's Horton the Elephant, Horton.


See also

* Animal track * Beehive fences use elephants' fear of bees to minimise conflict with humans * Desert elephant * Elephants' graveyard * List of individual elephants * Motty, captive hybrid of an Asian and African elephant * National Elephant Day (Thailand)


References


Bibliography

* ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** ** * * *


Further reading

* * Nance, Susan (2013). ''Entertaining Elephants: Animal Agency and the Business of the American Circus''. Baltimore, MD: Johns Hopkins University Press, 2013. * John Godfrey Saxe, Saxe, John Godfrey (1872). s:The Blindmen and the Elephant, "The Blindmen and the Elephant" at Wikisource. ''The Poems of John Godfrey Saxe''. *


External links


International Elephant Foundation
{{Featured article Elephants, Herbivorous mammals Tool-using mammals Articles containing video clips Mammal common names Pliocene first appearances