View source for Biology:Elephant

{{Short description|Large terrestrial mammals with trunks from Africa and Asia}}
{{Paraphyletic group
| name = Elephants
| image = Elephant Diversity.jpg
| image_alt = From top left to right: the [[Biology:African bush elephant|African bush elephant]], the [[Biology:Asian elephant|Asian elephant]] and [[Biology:African forest elephant|African forest elephant]].
| image_caption = From top left to right: the [[Biology:African bush elephant|African bush elephant]], the [[Biology:Asian elephant|Asian elephant]] and [[Biology:African forest elephant|African forest elephant]].
| auto = yes
| fossil_range = {{Fossil range|Pliocene|Present}}
| parent = Elephantinae
| includes = * ''[[Biology:African elephant|Loxodonta]]'' <small>Anonymous, 1827</small>
* ''[[Biology:Elephas|Elephas]]'' <small>[[Biography:Carl Linnaeus|Linnaeus]], [[Biology:10th edition of Systema Naturae|1758]]</small>
* {{extinct}}''[[Biology:Palaeoloxodon|Palaeoloxodon]]'' <small>Matsumoto, 1925</small>
| excludes = * {{extinct}}''[[Biology:Mammuthus|Mammuthus]]'' <small>Brookes, 1828</small>
* {{extinct}}''[[Biology:Primelephas|Primelephas]]'' <small>Maglio, 1970</small>
}}

'''Elephants''' are the largest existing land animals. Three living [[Biology:Species|species]] are currently recognised: the [[Biology:African bush elephant|African bush elephant]], the [[Biology:African forest elephant|African forest elephant]], and the [[Biology:Asian elephant|Asian elephant]]. They are an [[Biology:Paraphyly|informal grouping]] within the proboscidean family [[Biology:Elephantidae|Elephantidae]]. Elephantidae is the only surviving family of proboscideans; extinct members include the [[Biology:Mastodon|mastodon]]s. Elephantidae also contains several extinct groups, including the [[Biology:Mammoth|mammoth]]s and [[Biology:Straight-tusked elephant|straight-tusked elephant]]s. [[Biology:African elephant|African elephant]]s have larger ears and concave backs, whereas Asian elephants have smaller ears, and convex or level backs. Distinctive features of all elephants include a long [[Biology:Proboscis|proboscis]] called a trunk, tusks, large ear flaps, massive 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. The pillar-like legs carry their great weight.

Elephants are scattered throughout [[Place:Sub-Saharan Africa|sub-Saharan Africa]], [[Place:South Asia|South Asia]], and [[Place:Southeast Asia|Southeast Asia]] and are found in different habitats, including [[Earth:Savanna|savanna]]hs, forests, deserts, and [[Earth:Marsh|marsh]]es. They are herbivorous, and they stay near water when it is accessible. They are considered to be [[Biology:Keystone species|keystone species]], due to their impact on their environments.  Elephants have a [[Biology:Fission–fusion society|fission–fusion society]], 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 [[Social:Matriarchy|matriarch]].

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 [[Chemistry:Testosterone|testosterone]] and aggression known as musth, which helps them gain [[Dominance (ethology)|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 [[Biology:Elephant communication|communicate]] by touch, sight, smell, and sound; elephants use [[Physics:Infrasound|infrasound]], and [[Physics:Seismic communication|seismic communication]] over long distances. Elephant intelligence has been compared with that of [[Biology:Primate|primate]]s and [[Biology:Cetacea|cetacea]]ns. They appear to have [[Philosophy:Self-awareness|self-awareness]], and appear to show [[Philosophy:Empathy|empathy]] for dying and dead family members.

African bush elephants and Asian elephants are listed as [[Biology:Endangered species|endangered]] and African forest elephants as [[Biology:Critically Endangered|critically endangered]] by the [[Organization:International Union for Conservation of Nature|International Union for Conservation of Nature]] (IUCN). One of the biggest threats to elephant populations is the [[Finance:Ivory trade|ivory trade]], as the animals are [[Biology:Poaching|poached]] for their ivory tusks. Other threats to wild elephants include [[Earth:Habitat destruction|habitat destruction]] and conflicts with local people. Elephants are used as [[Biology:Working animal|working animal]]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 circuses. Elephants are highly recognisable and have been featured in art, folklore, religion, literature, and [[Social:Popular culture|popular culture]].

==Etymology==
The word "elephant" is based on the [[Social:Latin|Latin]] ''elephas'' (genitive ''elephantis'') ("elephant"), which is the [[Social:Latinisation of names|Latinised]] form of the [[Social:Greek language|Greek]] ἐλέφας (''elephas'') (genitive ἐλέφαντος (''elephantos''<ref name="LSJ">{{LSJ|e)le/fas|ἐλέφας|ref}}</ref>), probably from a non-[[Social:Indo-European languages|Indo-European]] language, likely [[Social:Phoenician language|Phoenician]].<ref name=etymology>{{cite dictionary|author=Harper, D|title=Elephant|dictionary=Online Etymology Dictionary|url=http://www.etymonline.com/index.php?term=elephant|access-date=25 October 2012}}</ref> It is attested in Mycenaean Greek as ''e-re-pa'' (genitive ''e-re-pa-to'') in [[Social:Linear B|Linear B]] syllabic script.<ref>{{Cite journal |url=https://www.academia.edu/2229199 |title=Ivory and horn production in Mycenaean texts |journal=Kosmos. Jewellery, Adornment and Textiles in the Aegean Bronze Age |last1=Lujan |first1=E. R. |last2=Bernabe |first2=A |access-date=22 January 2013}}</ref><ref>{{cite web|url=http://www.palaeolexicon.com/default.aspx?static=12&wid=189|title=elephant|publisher=Palaeolexicon, Word study tool of ancient languages|access-date=19 January 2013}}</ref> As in Mycenaean Greek, [[Biography:Homer|Homer]] used the Greek word to mean ivory, but after the time of Herodotus, it also referred to the animal.<ref name="LSJ" /> The word "elephant" appears in [[Social:Middle English|Middle English]] as ''olyfaunt'' (c.1300) and was borrowed from [[Social:Old French|Old French]] ''oliphant'' (12th century).<ref name=etymology />

==Taxonomy and phylogeny==
{{cladogram|style=font-size:70%|align=right|caption=A cladogram of the elephants within [[Biology:Afrotheria|Afrotheria]] based on molecular evidence<ref name=tabuce>{{cite journal|url=http://phylodiversity.net/azanne/csfar/images/d/d9/Afrotherian_mammals.pdf|first1=R.|last1=Tabuce| first2=R. J.|last2=Asher|first3=T.|last3=Lehmann|s2cid=46133294|year=2008|title=Afrotherian mammals: a review of current data| journal=Mammalia|volume=72|pages=2–14|doi=10.1515/MAMM.2008.004}}</ref>
|{{clade
|label1=[[Biology:Afrotheria|Afrotheria]]
|1={{Clade
   |label1=[[Biology:Afroinsectiphilia|Afroinsectiphilia]]
   |1={{Clade
      |label1=[[Biology:Tubulidentata|Tubulidentata]]
                 |1=[[Biology:Orycteropodidae|Orycteropodidae]] [[File:Aardvark2 (PSF) colourised.png|60 px]]
      |label2=[[Biology:Afroinsectivora|Afroinsectivora]]
      |2={{Clade
         |label1=[[Biology:Macroscelidea|Macroscelidea]]
                    |1=[[Biology:Macroscelididae|Macroscelididae]] [[File:Rhynchocyon chrysopygus-J Smit white background.jpg|50 px]]
         |label2=[[Biology:Afrosoricida|Afrosoricida]]
                     |2={{Clade
                         |1=[[Biology:Chrysochloridae|Chrysochloridae]] [[File:The animal kingdom, arranged according to its organization, serving as a foundation for the natural history of animals (Pl. 18) (Chrysochloris asiatica).jpg|50 px]]
                         |2=[[Biology:Tenrecidae|Tenrecidae]] [[File:Brehms Thierleben - Allgemeine Kunde des Thierreichs (1876) (Tenrec ecaudatus).jpg|50 px]]
      }} }} }}
   |label2=[[Biology:Paenungulata|Paenungulata]]
   |2={{Clade
      |label1=[[Biology:Hyrax|Hyracoidea]]
                 |1=[[Biology:Hyrax|Procaviidae]] [[File:DendrohyraxEminiSmit white background.jpg|50 px]]
      |label2=[[Biology:Tethytheria|Tethytheria]]
      |2={{Clade
         |label1=[[Biology:Proboscidea|Proboscidea]]
                    |1=[[Biology:Elephantidae|Elephantidae]] [[File:Elephas africanus - 1700-1880 - Print - Iconographia Zoologica - (white background).jpg|70 px]]
         |label2=[[Biology:Sirenia|Sirenia]]
                     |2={{Clade
                         |1=[[Biology:Dugongidae|Dugongidae]] [[File:Dugong dugon Hardwicke white background.jpg|60 px]]
                         |2=[[Biology:Trichechidae|Trichechidae]] [[File:Manatee white background.jpg|60 px]]
         }} }} }} }} }}}}
{{cladogram|style=font-size:70%|caption=Proboscidea phylogeny based on upper molars.<ref name=evolution />|align=right
|{{clade
|label1=[[Biology:Proboscidea|Proboscidea]]
|1={{clade
|1=early proboscideans, e.g. ''[[Biology:Moeritherium|Moeritherium]]'' [[File:Moeritherium NT small.jpg|70px]]
|2={{clade
   |1=[[Biology:Deinotheriidae|Deinotheriidae]] [[File:Deinotherium12.jpg|70px]]
   |label2=[[Biology:Elephantiformes|Elephantiformes]]
   |2={{clade
      |1=[[Biology:Mammutidae|Mammutidae]] [[File:BlankMastodon.jpg|70px]]
       |2={{clade
         |1=[[Biology:Gomphotheriidae|Gomphotheriidae]] [[File:Gomphotherium NT small.jpg|70px]]
         |2={{clade
            |1=[[Biology:Stegodontidae|Stegodontidae]] [[File:Stegodon Siwalik Hills.jpg|110px]]
            |label2=[[Biology:Elephantidae|Elephantidae]]
            |2={{clade
               |1=''[[Biology:Loxodonta|Loxodonta]]'' [[File:African Bush Elephant.jpg|50px]]
               |2={{clade
                  |1=''[[Biology:Mammuthus|Mammuthus]]'' [[File:Mammuthus trogontherii122DB.jpg|70px]]
                  |2=''[[Biology:Elephas|Elephas]]'' [[File:Elephas maximus (Bandipur).jpg|70px]]
                  }}
               }}
            }}
         }}
      }}
   }}
 }}
}}
}}
{{cladogram|style=font-size:70%|align=right|caption=Phylogeny of modern elephants and close extinct relatives based on molecular evidence<ref name="Palkopoulou"/>
|{{clade
|label1=Elephantimorpha
|1={{clade
   |label1=[[Biology:Elephantidae|Elephantidae]]
   |1={{clade
      |1={{clade
         |1={{clade
            |1=''Mammuthus primigenius'' [[File:Woolly mammoth model Royal BC Museum in Victoria.jpg|50px]]
            |2=''Mammuthus columbi'' [[File:Archidiskodon imperator121.jpg|50px]]
            }}
         |2=''Elephas maximus'' [[File:Elephas maximus (Bandipur).jpg|50px]]
         }}
      |2={{clade
         |1={{clade
            |1=''Loxodonta cyclotis'' [[File:African Forest Elephant.jpg|50px]]
            |2=''Palaeoloxodon antiquus'' [[File:Elephas-antiquus.jpg|50px]]
            }} 
         |2=''Loxodonta africana'' [[File:African Bush Elephant.jpg|50px]]
         }} 
      }}
   |2=''Mammut americanum'' [[File:BlankMastodon.jpg|50px]]
   }}
}}
}}
Elephants belong to the family [[Biology:Elephantidae|Elephantidae]], the sole remaining family within the order [[Biology:Proboscidea|Proboscidea]] which belongs to the superorder [[Biology:Afrotheria|Afrotheria]]. Their closest extant relatives are the [[Biology:Sirenia|sirenia]]ns ([[Biology:Dugong|dugong]]s and [[Biology:Manatee|manatee]]s) and the [[Biology:Hyrax|hyrax]]es, with which they share the [[Biology:Clade|clade]] [[Biology:Paenungulata|Paenungulata]] within the superorder [[Biology:Afrotheria|Afrotheria]].<ref name="Ozawa">{{cite journal|author1=Kellogg, M. |author2=Burkett, S. |author3=Dennis, T. R. |author4=Stone, G. |author5=Gray, B. A. |author6=McGuire, P. M. |author7=Zori, R. T. |author8=Stanyon, R. |year=2007|title=Chromosome painting in the manatee supports Afrotheria and Paenungulata|journal=Evolutionary Biology|volume=7|page=6|doi=10.1186/1471-2148-7-6 |pmid=17244368 |pmc=1784077}}</ref> Elephants and sirenians are further grouped in the clade [[Biology:Tethytheria|Tethytheria]].<ref name=Ozawa2>{{cite journal|author1=Ozawa, T. |author2=Hayashi, S. |author3=Mikhelson, V. M. |s2cid=417046 |year=1997|title=Phylogenetic position of mammoth and Steller's sea cow within tethytheria demonstrated by mitochondrial DNA sequences|journal=Journal of Molecular Evolution|volume=44|issue=4|pages=406–13|doi=10.1007/PL00006160|pmid=9089080|bibcode=1997JMolE..44..406O}}</ref>

Three species of elephants are recognised; the [[Biology:African bush elephant|African bush elephant]] (''[[Biology:Loxodonta|Loxodonta]] africana'') and [[Biology:African forest elephant|forest elephant]] (''Loxodonta cyclotis'') of [[Place:Sub-Saharan Africa|sub-Saharan Africa]], and the [[Biology:Asian elephant|Asian elephant]] (''[[Biology:Elephas|Elephas]] maximus'') of [[Place:South Asia|South]] and [[Place:Southeast Asia|Southeast Asia]].<ref name=MSW3>{{cite book|author=Shoshani, J.|year=2005|contribution=Order Proboscidea|editor1=Wilson, D. E. |editor2=Reeder, D. M |url=https://www.google.com/books?id=JgAMbNSt8ikC&pg=PA91#v=onepage|title=Mammal Species of the World: A Taxonomic and Geographic Reference|volume=1 |edition= 3rd|publisher=Johns Hopkins University Press|pages=90–91|isbn=978-0-8018-8221-0|oclc=62265494}}</ref> African elephants have larger ears, a concave back, more wrinkled skin, a sloping abdomen, and two finger-like extensions at the tip of the trunk. Asian elephants have smaller ears, a convex or level back, smoother skin, a horizontal abdomen that occasionally sags in the middle and one extension at the tip of the trunk. The looped ridges on the molars are narrower in the Asian elephant while those of the African are more diamond-shaped. The Asian elephant also has dorsal bumps on its head and some patches of depigmentation on its skin.<ref name=Shoshani38>Shoshani, pp. 38–41.</ref>

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 [[Place:West Africa|western]] and [[Place:Central Africa|Central Africa]].<ref name=Shoshani42 /> Both were traditionally considered a single species, ''Loxodonta africana'', but molecular studies have affirmed their status as separate species.<ref name=DNA>{{cite journal|author1=Rohland, N. |author2=Reich, D. |author3=Mallick, S. |author4=Meyer, M. |author5=Green, R. E. |author6=Georgiadis, N. J. |author7=Roca, A. L. |author8=Hofreiter, M. | title =Genomic DNA Sequences from Mastodon and Woolly Mammoth Reveal Deep Speciation of Forest and Savanna Elephants| journal = PLOS Biology | volume =8| issue =12| year =2010| doi=10.1371/journal.pbio.1000564| page =e1000564 | pmid =21203580 | pmc =3006346| editor1-last =Penny| editor1-first =David}}</ref><ref name="Ishida">{{cite journal|author1=Ishida, Y. |author2=Oleksyk, T. K. |author3=Georgiadis, N. J. |author4=David, V. A. |author5=Zhao, K. |author6=Stephens, R. M. |author7=Kolokotronis, S.-O. |author8=Roca, A. L. |year=2011|title=Reconciling apparent conflicts between mitochondrial and nuclear phylogenies in African elephants|journal=PLOS ONE|volume=6|issue=6|page=e20642|doi=10.1371/journal.pone.0020642|editor1-last=Murphy|editor1-first=William J|pmid=21701575|pmc=3110795|bibcode=2011PLoSO...620642I|doi-access=free }}</ref><ref name="genomic">{{Cite journal|title = Elephant Natural History: A Genomic Perspective|journal = Annual Review of Animal Biosciences|date = 2015|pmid = 25493538|pages = 139–167|volume = 3|issue = 1|doi = 10.1146/annurev-animal-022114-110838|first1 = Alfred L.|last1 = Roca|first2 = Yasuko|last2 = Ishida|first3 = Adam L.|last3 = Brandt|first4 = Neal R.|last4 = Benjamin|first5 = Kai|last5 = Zhao|first6 = Nicholas J.|last6 = Georgiadis}}</ref> In 2017, DNA sequence analysis showed that ''L. cyclotis'' is more closely related to the extinct ''Palaeoloxodon antiquus'', than it is to ''L. africana,'' possibly undermining the genus ''Loxodonta'' as a whole.<ref>{{cite journal|last1=Meyer|first1=Matthias|title=Palaeogenomes of Eurasian straight-tusked elephants challenge the current view of elephant evolution|journal=eLife|date=2017|doi=10.7554/eLife.25413|pmid=28585920|pmc=5461109|volume=6}}</ref>

===Evolution and extinct relatives===
Over 180 extinct members and three major evolutionary radiations of the order Proboscidea have been recorded.<ref>{{cite book|last1=Kingdon|first1=Jonathan|title=Mammals of Africa|date=2013|publisher=Bloomsbury|page=173|url=https://books.google.com/books?id=B_07noCPc4kC|isbn=9781408189962}}</ref> The earliest proboscids, the African ''[[Biology:Eritherium|Eritherium]]'' and ''[[Biology:Phosphatherium|Phosphatherium]]'' of the late [[Earth:Paleocene|Paleocene]], heralded the first radiation.<ref name="Gheerbrant">{{cite journal |author=Gheerbrant, E. |year=2009 |title=Paleocene emergence of elephant relatives and the rapid radiation of African ungulates |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=106 |issue=26 |pages=10717–10721 |doi=10.1073/pnas.0900251106 |pmid=19549873 |pmc=2705600|bibcode=2009PNAS..10610717G |doi-access=free }}</ref> The [[Earth:Eocene|Eocene]] included ''[[Biology:Numidotherium|Numidotherium]]'', ''[[Biology:Moeritherium|Moeritherium]],'' and ''[[Biology:Barytherium|Barytherium]]'' from Africa. These animals were relatively small and aquatic. Later on, genera such as ''[[Biology:Phiomia|Phiomia]]'' and ''[[Biology:Palaeomastodon|Palaeomastodon]]'' arose; the latter likely inhabited forests and open woodlands. Proboscidean diversity declined during the Oligocene.<ref>Sukumar, pp. 13–16.</ref> One notable species of this epoch was ''[[Biology:Eritreum|Eritreum]] melakeghebrekristosi'' of the [[Place:Horn of Africa|Horn of Africa]], which may have been an ancestor to several later species.<ref name=link>{{cite journal|author1=Shoshani, J. |author2=Walter, R. C. |author3=Abraha, M. |author4=Berhe, S. |author5=Tassy, P. |author6=Sanders, W. J. |author7=Marchant, G. H. |author8=Libsekal, Y. |author9=Ghirmai, T. |author10=Zinner, D. |year=2006|title=A proboscidean from the late Oligocene of Eritrea, a "missing link" between early Elephantiformes and Elephantimorpha, and biogeographic implications|journal=Proceedings of the National Academy of Sciences|volume=103|issue=46|pages=17296–301|doi=10.1073/pnas.0603689103|bibcode=2006PNAS..10317296S |pmid=17085582 |pmc=1859925|doi-access=free }}</ref> The beginning of the Miocene saw the second diversification, with the appearance of the [[Biology:Deinotheriidae|deinotheres]] and the [[Biology:Mammutidae|mammutids]]. The former were related to ''Barytherium'' and lived in Africa and Eurasia,<ref name=Sukumar17>Sukumar, pp. 16–19.</ref> while the latter may have descended from ''Eritreum''<ref name=link /> and spread to North America.<ref name=Sukumar17 />

The second radiation was represented by the emergence of the [[Biology:Gomphothere|gomphothere]]s in the Miocene,<ref name=Sukumar17 /> which likely evolved from ''Eritreum''<ref name=link /> and originated in Africa, spreading to every continent except Australia and Antarctica. Members of this group included ''[[Biology:Gomphotherium|Gomphotherium]]'' and ''[[Biology:Platybelodon|Platybelodon]]''.<ref name=Sukumar17 /> The third radiation started in the late Miocene and led to the arrival of the elephantids, which descended from, and slowly replaced, the gomphotheres.<ref>Sukumar, p. 22.</ref> The African ''[[Biology:Primelephas|Primelephas]] gomphotheroides'' gave rise to ''Loxodonta'', ''Mammuthus,'' and ''Elephas''. ''Loxodonta'' branched off earliest around the Miocene and Pliocene boundary while ''Mammuthus'' and ''Elephas'' diverged later during the early Pliocene. ''Loxodonta'' remained in Africa while ''Mammuthus'' and ''Elephas'' spread to Eurasia, and the former reached North America. At the same time, the [[Biology:Stegodontidae|stegodontids]], another proboscidean group descended from gomphotheres, spread throughout Asia, including the Indian subcontinent, China, southeast Asia, and Japan. Mammutids continued to evolve into new species, such as the [[Biology:Mastodon|American mastodon]].<ref>Sukumar, pp. 24–27.</ref>

At the beginning of the [[Earth:Pleistocene|Pleistocene]], elephantids experienced a high rate of [[Biology:Speciation|speciation]].<ref name=Sukmar44/> The Pleistocene also saw the arrival of ''[[Biology:Palaeoloxodon namadicus|Palaeoloxodon namadicus]]'', the largest terrestrial mammal of all time.<ref name="Larramendi, A. 2015">{{cite journal | author = Larramendi A | year = 2015 | title = Shoulder height, body mass and shape of proboscideans | journal = Acta Palaeontologica Polonica | doi = 10.4202/app.00136.2014 | doi-access = free }}</ref> ''[[Biology:Loxodonta atlantica|Loxodonta atlantica]]'' became the most common species in northern and southern Africa but was replaced by ''Elephas iolensis'' later in the Pleistocene. Only when ''Elephas'' disappeared from Africa did ''Loxodonta'' become dominant once again, this time in the form of the modern species. ''Elephas'' diversified into new species in Asia, such as ''E.&nbsp;hysudricus'' and ''E.&nbsp;platycephus'';<ref name=Sukumar28 /> the latter the likely ancestor of the modern Asian elephant.<ref name=Sukmar44>Sukumar, p. 44.</ref> ''Mammuthus'' evolved into several species, including the well-known [[Biology:Woolly mammoth|woolly mammoth]].<ref name=Sukumar28>Sukumar, pp. 28–31.</ref> 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''.<ref name="Palkopoulou">{{cite journal|author=Palkopoulou, E.|display-authors=0|year=2018|title=A comprehensive genomic history of extinct and living elephants|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=115|issue=11|pages=E2566–E2574|doi=10.1073/pnas.1720554115|pmid=29483247|pmc=5856550|doi-access=free}}</ref> In the [[Earth:Late Pleistocene|Late Pleistocene]], most proboscidean species vanished during the [[Earth:Quaternary glaciation|Quaternary glaciation]] which [[Earth:Quaternary extinction event|killed off]] 50% of genera weighing over {{convert|5|kg|lb|abbr=on}} worldwide.<ref>Sukumar, pp. 36–37.</ref>

Proboscideans experienced several evolutionary trends, such as an increase in size, which led to many giant species that stood up to {{convert|500|cm|ftin|0|abbr=on}} tall.<ref name="Larramendi, A. 2015"/> As with other [[Earth:Megaherbivore|megaherbivore]]s, including the extinct [[Biology:Sauropoda|sauropod]] [[Biology:Dinosaur|dinosaur]]s, the large size of elephants likely developed to allow them to survive on vegetation with low nutritional value.<ref>{{cite book|author=Carpenter, K.|year=2006 |chapter=Biggest of the big: a critical re-evaluation of the mega-sauropod ''Amphicoelias fragillimus'' Cope, 1878 |editor=Foster, J.R. |editor2=Lucas, S.G. |title= Paleontology and Geology of the Upper Jurassic Morrison Formation |series=New Mexico Museum of Natural History and Science Bulletin |publisher=New Mexico Museum of Natural History and Science |volume=36 |pages=131–138}}</ref> Their limbs grew longer and the feet shorter and broader.<ref name=evolution /> The feet were originally [[Biology:Plantigrade|plantigrade]] and developed into a [[Biology:Digitigrade|digitigrade]] stance with cushion pads and the [[Biology:Sesamoid bone|sesamoid bone]] providing support.<ref>{{cite journal|author1=Hutchinson, J. R. |author2=Delmer, C. |author3=Miller, C. E. |author4=Hildebrandt, T. |author5=Pitsillides, A. A. |author6=Boyde, A. |s2cid=206536505 |year=2011|title=From flat foot to fat foot: structure, ontogeny, function, and evolution of elephant 'sixth toes' |journal=Science |volume=334 |issue=6063 |pages=1699–1703 |doi=10.1126/science.1211437 |pmid=22194576 |bibcode=2011Sci...334R1699H}}</ref> Early proboscideans developed longer mandibles and smaller [[Biology:Skull|craniums]] while more derived ones developed shorter mandibles, which shifted the head's [[Physics:Center of mass|centre of gravity]]. The skull grew larger, especially the cranium, while the neck shortened to provide better support for the skull. The increase in size led to the development and elongation of the mobile trunk to provide reach. The number of premolars, incisors and [[Biology:Canine tooth|canines]] decreased.<ref name=evolution />

The cheek teeth (molars and premolars) of proboscideans became larger and more specialized, especially after elephants started to switch from [[Biology:C3 carbon fixation|C3-plants]] to [[Biology:C4 carbon fixation|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.<ref>{{Cite news|url=http://www.thehindu.com/sci-tech/energy-and-environment/the-dental-factor-in-elephant-evolution/article5093769.ece|title=The dental factor in elephant evolution|author=K.S. Rajgopal|newspaper=The Hindu|date=4 September 2013}}</ref><ref>{{cite web|url=http://www.livescience.com/13550-grazing-animals-shaped-human-evolution.html|title=Humanity's Grassroots: How Grazing Animals Shaped Evolution|work=LiveScience.com|date=4 April 2011}}</ref> 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.<ref name=evolution>{{cite journal|author=Shoshani, J. |year=1998 |title=Understanding proboscidean evolution: a formidable task|journal=Trends in Ecology and Evolution|volume=13|issue=12|pages=480–87|doi=10.1016/S0169-5347(98)01491-8|pmid=21238404}}</ref> Elephants retain certain features from their aquatic ancestry, such as their [[Medicine:Middle ear|middle ear]] anatomy.<ref name=snorkel />

Several species of proboscideans lived on islands and experienced [[Biology:Insular dwarfism|insular dwarfism]]. This occurred primarily during the Pleistocene when some elephant populations became isolated by fluctuating sea levels, although dwarf elephants did exist earlier in the Pliocene. These elephants likely grew smaller on islands due to a lack of large or viable predator populations and limited resources. By contrast, small mammals such as [[Biology:Rodent|rodent]]s develop [[Biology:Island gigantism|gigantism]] in these conditions. [[Biology:Dwarf elephant|Dwarf elephant]]s are known to have lived in Indonesia, the Channel Islands of California, and several islands of the [[Place:Mediterranean Sea|Mediterranean]].<ref name=Sukumar31>Sukumar, pp. 31–33.</ref>

== Anatomy and morphology ==
===Size===
[[File:Elephant skeleton.jpg|thumb|right|African bush elephant skeleton]]
Elephants are the largest living terrestrial animals. African bush elephants are the largest species, with males being {{cvt|304|-|336|cm|ftin|0|abbr=on}} tall at the shoulder with a body mass of {{cvt|5.2|-|6.9|MT|ST|1|abbr=on}} and females standing {{cvt|247|-|273|cm|ftin|0|abbr=on}} tall at the shoulder with a body mass of {{cvt|2.6|-|3.5|MT|ST|1|abbr=on}}. Male Asian elephants are usually about {{cvt|261|-|289|cm|ftin|0|abbr=on}} tall at the shoulder and {{cvt|3.5|-|4.6|MT|ST|1|abbr=on}} whereas females are {{cvt|228|-|252|cm|ftin|0|abbr=on}} tall at the shoulder and {{cvt|2.3|-|3.1|MT|ST|1|abbr=on}}. African forest elephants are the smallest species, with males usually being around {{cvt|209|-|231|cm|ftin|0|abbr=on}} tall at the shoulder and {{cvt|1.7|-|2.3|MT|ST|1|abbr=on}}. Male African bush elephants are typically 23% taller than females, whereas male Asian elephants are only around 15% taller than females.<ref name="Larramendi, A. 2015"/>

===Bones===
The skeleton of the elephant is made up of 326–351 bones.<ref name=Shoshani68 /> 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.<ref>{{cite web|author=Somgrid, C.|title=Elephant Anatomy and Biology: Skeletal system|publisher=Elephant Research and Education Center, Department of Companion Animal and Wildlife Clinics, Faculty of Veterinary Medicine, Chiang Mai University|url=http://www.asianelephantresearch.com/about-elephant-anatomy-and-biology-p1.php#skeleton|access-date=21 September 2012|archive-url=https://web.archive.org/web/20120613191055/http://www.asianelephantresearch.com/about-elephant-anatomy-and-biology-p1.php#skeleton|archive-date=13 June 2012|url-status=dead}}</ref>

===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 in every direction.<ref>Kingdon, p. 11.</ref> The skull contains air cavities ([[Biology:Sinus (anatomy)|sinuses]]) that reduce the weight of the skull while maintaining overall strength. These cavities give the inside of the skull a honeycomb-like appearance. The cranium is particularly large and provides enough room for the attachment of muscles to support the entire head. The lower jaw is solid and heavy.<ref name=Shoshani68>Shoshani, pp. 68–70.</ref> Because of the size of the head, the neck is relatively short to provide better support.<ref name=evolution /> Lacking a [[Medicine:Lacrimal apparatus|lacrimal apparatus]], the eye relies on the [[Biology:Harderian gland|harderian gland]] to keep it moist. A durable [[Biology:Nictitating membrane|nictitating membrane]] protects the eye globe. The animal's [[Physics:Field of view|field of vision]] is compromised by the location and limited mobility of the eyes.<ref name="sense">{{cite web|author=Somgrid, C.|title=Elephant Anatomy and Biology: Special sense organs|publisher=Elephant Research and Education Center, Department of Companion Animal and Wildlife Clinics, Faculty of Veterinary Medicine, Chiang Mai University|url=http://www.asianelephantresearch.com/about-elephant-anatomy-and-biology-p4.php#Special|access-date=21 September 2012|archive-url=https://web.archive.org/web/20130729045534/http://www.asianelephantresearch.com/about-elephant-anatomy-and-biology-p4.php#Special|archive-date=29 July 2013|url-status=dead}}</ref> Elephants are considered [[Medicine:Dichromacy|dichromats]]<ref>{{cite journal|author1=Yokoyama, S. |author2=Takenaka, N. |author3=Agnew, D. W. |author4=Shoshani, J. |year=2005|title=Elephants and human color-blind deuteranopes have identical sets of visual pigments|journal=Genetics|volume=170|issue=1|pages=335–44|doi=10.1534/genetics.104.039511|pmid=15781694|pmc=1449733}}</ref> and they can see well in dim light but not in bright light.<ref name=cognition />
[[File:Angry elephant ears.jpg|thumb|African bush elephant with ears spread in a threat or attentive position; note the visible blood vessels]]

=== Ears ===
Elephant ears have thick bases with thin tips. The ear flaps, or pinnae, contain numerous blood vessels called [[Biology:Capillary|capillaries]]. Warm blood flows into the capillaries, helping to release excess body heat into the environment. This occurs when the pinnae are still, and the animal can enhance the effect by flapping them. Larger ear surfaces contain more capillaries, and more heat can be released. Of all the elephants, African bush elephants live in the hottest climates, and have the largest ear flaps.<ref>{{cite journal|author=Narasimhan, A.|s2cid=121443269|year=2008|title=Why do elephants have big ear flaps?|journal=Resonance|volume=13|issue=7|pages=638–47|doi=10.1007/s12045-008-0070-5}}</ref> Elephants are capable of hearing at low frequencies and are most sensitive at 1 [[Hertz|kHz]] (in close proximity to the Soprano C).<ref>{{cite journal|author1=Reuter, T.|author2=Nummela, S.|author3=Hemilä, S.|year=1998|title=Elephant hearing|journal=Journal of the Acoustical Society of America|volume=104|issue=2|pages=1122–23|url=http://roadecology.ucdavis.edu/%5C/pdflib/Winter2005/Rodwell_Reuter_ele_ear.pdf|doi=10.1121/1.423341|pmid=9714930|bibcode=1998ASAJ..104.1122R|url-status=dead|archive-url=https://web.archive.org/web/20121207065356/http://roadecology.ucdavis.edu/pdflib/Winter2005/Rodwell_Reuter_ele_ear.pdf|archive-date=7 December 2012|df=dmy-all}}</ref>

=== Trunk ===
{{multiple image |align=right |direction=vertical
|image1=African elephant warning raised trunk.jpg|caption1=African bush elephant with its trunk raised, a behaviour often adopted when trumpeting
|image2=Asian Elephant, Royal Chitwan National Park.jpg|caption2=Asian elephant drinking water with trunk
}}
The trunk, or [[Biology:Proboscis|proboscis]], is a fusion of the nose and upper lip, although in early [[Biology:Fetus|fetal]] life, the upper lip and trunk are separated.<ref name=evolution /> The trunk is elongated and specialised to become the elephant's most important and versatile appendage. It contains up to 150,000 separate [[Biology:Muscle fascicle|muscle fascicle]]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 [[Biology:Transverse plane|transverse]] and radiating muscles. The muscles of the trunk connect to a bony opening in the skull. The nasal septum is composed of tiny muscle units that stretch horizontally between the nostrils. [[Biology:Cartilage|Cartilage]] divides the nostrils at the base.<ref name=Shoshani74>Shoshani, pp. 74–77.</ref> As a [[Biology:Muscular hydrostat|muscular hydrostat]], the trunk moves by precisely coordinated muscle contractions. The muscles work both with and against each other. A unique proboscis nerve – formed by the maxillary and [[Biology:Facial nerve|facial nerve]]s – runs along both sides of the trunk.<ref name=trunk />

Elephant trunks have multiple functions, including breathing, [[Biology:Olfaction|olfaction]], touching, grasping, and sound production.<ref name=evolution /> The animal's sense of smell may be four times as sensitive as that of a bloodhound.<ref name=Sukumar149 /> The trunk's ability to make powerful twisting and coiling movements allows it to collect food, wrestle with other elephants,<ref name=Kingdon9>Kingdon, p. 9.</ref> and lift up to {{convert|350|kg|lb|abbr=on}}.<ref name=evolution /> It can be used for delicate tasks, such as wiping an eye and checking an orifice,<ref name=Kingdon9 /> and is capable of cracking a peanut shell without breaking the seed.<ref name=evolution /> With its trunk, an elephant can reach items at heights of up to {{convert|7|m|ft|abbr=on}} and dig for water under mud or sand.<ref name=Kingdon9 /> Individuals may show lateral preference when grasping with their trunks: some prefer to twist them to the left, others to the right.<ref name="trunk">{{cite journal|author1=Martin, F.|author2=Niemitz C. |year=2003|title="Right-trunkers" and "left-trunkers": side preferences of trunk movements in wild Asian elephants (''Elephas maximus'')|journal=Journal of Comparative Psychology|volume=117|issue=4|pages=371–79|doi=10.1037/0735-7036.117.4.371|pmid=14717638}}</ref> Elephants are capable of dilating their nostrils at a radius of nearly 30%, increasing the nasal volume by 64%, and can inhale at over {{convert|150|m/s|abbr=on}} which is around 30 times the speed of a human sneeze.<ref name="Schulz">{{cite journal|author=Schulz, A. K.; Ning Wu, Jia; Sara Ha, S. Y.; Kim, G.|year=2021|title=Suction feeding by elephants|journal=Journal of the Royal Society Interface|volume=18|issue=179|doi=10.1098/rsif.2021.0215|pmid=34062103|pmc=8169210|doi-access=free|pmc-embargo-date=June 2, 2022}}</ref> Elephants can suck up food and water both to spray in the mouth and, in the case of the later, to sprinkle on their bodies.<ref name=evolution /><ref name="Schulz"/> An adult Asian elephant is capable of holding {{convert|8.5|L|gal|abbr=on}} of water in its trunk.<ref name=Shoshani74 />  They will also spray dust or grass on themselves.<ref name=evolution /> When underwater, the elephant uses its trunk as a [[Engineering:Submarine snorkel|snorkel]].<ref name=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.<ref name=Shoshani38 /> Asian elephants have more muscle coordination and can perform more complex tasks.<ref name=Shoshani74 /> Losing the trunk would be detrimental to an elephant's survival,<ref name=evolution /> 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.<ref name=Shoshani74 /> Floppy trunk syndrome is a condition of trunk [[Engineering:Paralysis|paralysis]] in African bush elephants caused by the degradation of the [[Biology:Peripheral nervous system|peripheral nerves]] and muscles beginning at the tip.<ref name="NS">{{cite magazine|url=https://www.newscientist.com/article/mg13618470.700-lead-in-lake-blamed-for-floppy-trunks-.html|title=Lead in lake blamed for floppy trunks|author=Cole, M.|date=14 November 1992|magazine=New Scientist|access-date=25 June 2009}}</ref>

=== Teeth ===
{{multiple image |align=right |direction=vertical
|image1=Jaw of a deceased Loxodonta africana juvenile individual found within the Voyager Ziwani Safari Camp, on the edge of the Tsavo West National Park, near Ziwani, Kenya 3 (edited).jpg|caption1=Closeup of the cheek teeth of a dead juvenile bush elephant
|image2=2010-kabini-tusker-bark.jpg|caption2=Asian elephant eating tree bark, using its tusks to peel it off.
}}
Elephants usually have 26 teeth: the incisors, known as the [[Biology:Tusk|tusk]]s, 12 deciduous premolars, and 12 molars. Unlike most mammals, which grow baby teeth and then replace them with a single permanent set of adult teeth, elephants are [[Biology:Polyphyodont|polyphyodont]]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.<ref name=Shoshani70>Shoshani, pp. 70–71.</ref>

==== Tusks ====
The tusks of an elephant are modified second incisors in the upper jaw. They replace deciduous milk teeth at 6–12 months of age and grow continuously at about {{convert|17|cm|in|0|abbr=on}} a year. A newly developed tusk has a smooth [[Biology:Tooth enamel|enamel]] cap that eventually wears off. The [[Biology:Dentin|dentin]]e is known as ivory and its [[Cross section (geometry)|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 [[Chemistry:Calcite|calcite]]. 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 [[Biology:Pulp (tooth)|pulp]] 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.<ref name=Shoshani71 />

Like humans, who are typically [[Physics:Handedness|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 {{convert|300|cm|ftin|0|abbr=on}},<ref name=Shoshani71 /> but those of males tend to be thicker.<ref>Sukumar, p. 120</ref> In earlier times, elephant tusks weighing over 200 pounds (more than 90&nbsp;kg) were not uncommon, though it is rare today to see any over {{convert|100|lb|kg}}.<ref>"Still Life" by Bryan Christy. ''National Geographic Magazine'', August 2015, pp. 97, 104.</ref>

In the Asian species, only the males have large tusks. Female Asians have very small tusks, or none at all.<ref name=Shoshani71>Shoshani, pp. 71–74.</ref> Tuskless males exist and are particularly common among Sri Lankan elephants.<ref>{{cite book|author=Clutton-Brock, J.|year=1986|title=A Natural History of Domesticated Mammals|publisher=British Museum (Natural History)|page=208|isbn=978-0-521-34697-9}}</ref> Asian males can have tusks as long as Africans', but they are usually slimmer and lighter; the largest recorded was {{convert|302|cm|ftin|0|abbr=on}} long and weighed {{convert|39|kg|lb|0|abbr=on}}. Hunting for elephant ivory in Africa<ref>{{cite web|title=Elephants Evolve Smaller Tusks Due to Poaching|date=20 January 2008|publisher=Environmental News Network|url=http://www.enn.com/wildlife/article/29620|access-date=25 September 2012}}</ref> and Asia<ref>{{Cite web|date=2018-11-09|title=Under poaching pressure, elephants are evolving to lose their tusks|url=https://www.nationalgeographic.com/animals/article/wildlife-watch-news-tuskless-elephants-behavior-change|url-status=live|access-date=2021-10-28|website=National Geographic|language=en}}</ref> has led to [[Biology:Natural selection|natural selection]] for shorter tusks<ref>{{cite news|url=https://www.telegraph.co.uk/science/science-news/3322455/Why-elephants-are-not-so-long-in-the-tusk.html|archive-url=https://web.archive.org/web/20091018192954/http://www.telegraph.co.uk/science/science-news/3322455/Why-elephants-are-not-so-long-in-the-tusk.html|url-status=dead|archive-date=18 October 2009|title=Why elephants are not so long in the tusk|last=Gray|first=R.|date=20 January 2008|work=The Daily Telegraph|access-date=27 January 2013}}</ref><ref>{{cite journal|author1=Chiyo, P. I. |author2=Obanda, V. |author3=Korir, D. K. |title=Illegal tusk harvest and the decline of tusk size in the African elephant|journal=Ecology and Evolution|year=2015|volume=5|issue=22|pages=5216–5229|doi=10.1002/ece3.1769|pmid=30151125|pmc=6102531}}</ref> and tusklessness.<ref>{{cite journal|author1=Jachmann, H. |author2=Berry, P. S. M. |author3=Imae, H. |year=1995|title=Tusklessness in African elephants: a future trend|journal=African Journal of Ecology|volume=33|issue=3|pages=230–235|doi=10.1111/j.1365-2028.1995.tb00800.x}}</ref><ref>{{cite journal|author1=Kurt, F. |author2=Hartl, G. |author3=Tiedemann, R. |year=1995|title=Tuskless bulls in Asian elephant ''Elephas maximus''. History and population genetics of a man-made phenomenon.|journal=Acta Theriol.|volume=40|pages=125–144|doi=10.4098/at.arch.95-51|doi-access=free}}</ref>

=== Skin ===
[[File:Forest elephant group 4 (6987537249).jpg|thumb|An African forest elephant covering its skin with mud]]
An elephant's skin is generally very tough, at {{convert|2.5|cm|in|0|abbr=on}} thick on the back and parts of the head. The skin around the mouth, [[Biology:Anus|anus]], and inside of the ear is considerably thinner. Elephants typically have grey skin, but African elephants look brown or reddish after wallowing in coloured mud. Asian elephants have some patches of depigmentation, particularly on the forehead and ears and the areas around them. Calves have brownish or reddish hair, especially on the head and back. As elephants mature, their hair darkens and becomes sparser, but dense concentrations of hair and bristles remain on the end of the tail as well as the chin, [[Biology:Sex organ|genitals]] and the areas around the eyes and ear openings. Normally the skin of an Asian elephant is covered with more hair than its African counterpart.<ref name=Shoshani66>Shoshani, pp. 66–67.</ref> Their hair is thought to be for [[Physics:Thermoregulation|thermoregulation]], helping them lose heat in their hot environments.<ref name="Elephanthair">{{cite journal |last1=Myhrvold |first1=C. L. |last2=Stone |first2=H. A. |last3=Bou-Zeid |first3=E. |title=What Is the Use of Elephant Hair? |journal=PLOS ONE |date=10 October 2012 |volume=7 |issue=10 |pages=e47018 |doi=10.1371/journal.pone.0047018|pmid=23071700 |pmc=3468452 |bibcode=2012PLoSO...747018M |doi-access=free }}</ref>

An elephant uses mud as a sunscreen, protecting its skin from [[Physics:Ultraviolet|ultraviolet]] light. Although tough, an elephant's skin is very sensitive. Without regular [[Chemistry:Mud bath|mud bath]]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 [[Biology:Surface-area-to-volume ratio|surface-area-to-volume ratio]], which is many times smaller than that of a human. They have even been observed lifting up their legs, presumably in an effort to expose their soles to the air.<ref name=Shoshani66 />

=== Legs, locomotion, and posture ===
[[File:Elephant Walking animated.gif|thumb|An Asian elephant walking]]
To support the animal's weight, an elephant's limbs are positioned more vertically under the body than in most other mammals. The long bones of the limbs have cancellous bone in place of [[Biology:Medullary cavity|medullary cavities]]. This strengthens the bones while still allowing [[Biology:Haematopoiesis|haematopoiesis]].<ref name=Shoshani69>Shoshani, pp. 69–70.</ref> Both the front and hind limbs can support an elephant's weight, although 60% is borne by the front.<ref name="cushion">{{cite journal|author1=Weissengruber, G. E. |author2=Egger, G. F. |author3=Hutchinson, J. R. |author4=Groenewald, H. B. |author5=Elsässer, L. |author6=Famini, D. |author7=Forstenpointner, G. |year=2006|title=The structure of the cushions in the feet of African elephants (''Loxodonta africana'')|journal=Journal of Anatomy|volume=209|issue=6|pages=781–92|doi=10.1111/j.1469-7580.2006.00648.x|pmid=17118065|pmc=2048995}}</ref> 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 [[Biology:Ulna|ulna]] and [[Biology:Radius (bone)|radius]] are fixed in pronation; the "palm" of the manus faces backward.<ref name=Shoshani69 /> The pronator quadratus and the pronator teres are either reduced or absent.<ref>Shoshani, p. 74.</ref> The circular feet of an elephant have soft tissues or "cushion pads" beneath the [[Biology:Manus (anatomy)|manus]] or [[Biology:Pes (anatomy)|pes]], which distribute the weight of the animal.<ref name=cushion /> They appear to have a [[Biology:Sesamoid bone|sesamoid]], an extra "toe" similar in placement to a [[Biology:Giant panda|giant panda]]'s extra "thumb", that also helps in weight distribution.<ref>{{cite web |access-date=23 December 2011 |url=http://news.sciencemag.org/sciencenow/2011/12/elephants-have-a-sixth-toe.html? |title=Elephants Have a Sixth 'Toe' |publisher=ScienceMag.org |author=Pennisi, E. |date=22 December 2011 |url-status=dead |archive-url=https://web.archive.org/web/20120113093919/http://news.sciencemag.org/sciencenow/2011/12/elephants-have-a-sixth-toe.html |archive-date=13 January 2012 |df=dmy-all }}</ref> As many as five toenails can be found on both the front and hind feet.<ref name=Shoshani38 />

Elephants can move both forwards and backwards, but cannot trot, [[Biology:Jumping|jump]], or gallop. They use only two gaits when moving on land: the walk and a faster gait similar to running.<ref name=Shoshani69 /> 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.<ref name="kinetics">{{cite journal |author1 = Hutchinson, J. R.| author2 = Schwerda, D.| author3 = Famini, D. J.| author4 = Dale, R. H.| author5 = Fischer, M. S.| author6 = Kram, R.|year=2006|title=The locomotor kinematics of Asian and African elephants: changes with speed and size|journal=Journal of Experimental Biology|volume=209|issue=19|pages=3812–27|pmid=16985198 |doi = 10.1242/jeb.02443|doi-access=free}}</ref> Fast-moving elephants appear to 'run' with their front legs, but 'walk' with their hind legs and can reach a top speed of {{convert|25|km/h|abbr=on}}.<ref name="ReferenceA">{{cite journal|author1=Hutchinson, J. R. |author2=Famini, D. |author3=Lair, R. |author4=Kram, R. |s2cid=4403723 |year = 2003 | title = Biomechanics: Are fast-moving elephants really running? | journal = Nature | volume = 422 | pages = 493–94 | doi = 10.1038/422493a | pmid = 12673241 | issue = 6931|bibcode = 2003Natur.422..493H}}</ref> At this speed, most other [[Quadrupedalism|quadrupeds]] are well into a gallop, even accounting for leg length. Spring-like kinetics could explain the difference between the motion of elephants and other animals.<ref name="ReferenceA"/> During locomotion, the cushion pads expand and contract, and reduce both the pain and noise that would come from a very heavy animal moving.<ref name=cushion /> Elephants are capable swimmers. They have been recorded swimming for up to six hours without touching the bottom, and have travelled as far as {{convert|48|km|mi|abbr=on}} at a stretch and at speeds of up to {{convert|2.1|km/h|0|abbr=on}}.<ref>Shoshani, p. 60.</ref>

=== Organs ===
[[File:Em - Loxodonta africana heart - GMZ 2.jpg|thumb|right|African elephant heart in a jar]]
The brain of an elephant weighs {{convert|4.5|–|5.5|kg|lb|0|abbr=on}} compared to {{convert|1.6|kg|lb|0|abbr=on}} for a human brain. While the elephant brain is larger overall, it is proportionally smaller. At birth, an elephant's brain already weighs 30–40% of its adult weight. The [[Biology:Cerebrum|cerebrum]] and [[Biology:Cerebellum|cerebellum]] are well developed, and the [[Biology:Temporal lobe|temporal lobe]]s are so large that they bulge out laterally.<ref name=Shoshani78 /> The throat of an elephant appears to contain a pouch where it can store water for later use.<ref name=evolution /> The [[Medicine:Larynx|larynx]] of the elephant is the largest known among mammals. The vocal folds are long and are attached close to the [[Biology:Epiglottis|epiglottis]] 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.<ref name="vibratorypatterns">{{cite journal|author1= Herbest, C. T.| author2 = Švec, J. G.| author3 = Lohscheller, J.| author4 = Frey, R.| author5 = Gumpenberger, M.| author6 = Stoeger, A.| author7 = Fitch, W. T.|year=2013|title= Complex Vibratory Patterns in an Elephant Larynx|journal= Journal of Experimental Biology|volume=216|issue=21|pages=4054–4064|doi=10.1242/jeb.091009
|pmid=24133151|doi-access=free}}</ref>

The heart of an elephant weighs {{convert|12|–|21|kg|lb|abbr=on}}. It has a double-pointed [[Biology:Heart#Location and shape|apex]], an unusual trait among mammals.<ref name=Shoshani78 /> In addition, the [[Medicine:Ventricle (heart)|ventricles]] separate near the top of the heart, a trait they share with sirenians.<ref name="Marshall Cavendish">{{cite book| author=Anon|title=Mammal Anatomy: An Illustrated Guide |year=2010 |publisher=Marshall Cavendish |isbn=978-0-7614-7882-9 |pages=59}}</ref> 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.<ref>{{cite journal|author1=Benedict, F. G. |author2=Lee, R. C. |year=1936|title=The heart rate of the elephant|journal=Proceedings of the American Philosophical Society|volume=76|issue=3|pages=335–41|jstor=984548}}</ref> The blood vessels in most of the body are wide and thick and can withstand high blood pressures.<ref name="Marshall Cavendish"/> The lungs are attached to the diaphragm, and breathing relies mainly on the diaphragm rather than the expansion of the ribcage.<ref name=Shoshani78>Shoshani, pp. 78–79.</ref> Connective tissue exists in place of the pleural cavity. This may allow the animal to deal with the pressure differences when its body is underwater and its trunk is breaking the surface for air,<ref name="snorkel">{{cite journal|author=West, J. B.|s2cid=27321751|year=2002|title=Why doesn't the elephant have a pleural space?|journal=Physiology|volume=17|issue=2|pages=47–50|pmid=11909991|doi=10.1152/nips.01374.2001}}</ref> although this explanation has been questioned.<ref>{{cite news|title=How elephants 'snorkel'|work=BBC News|date=31 August 2002|access-date=3 November 2012|url=http://news.bbc.co.uk/2/hi/science/nature/2225846.stm}}</ref> Another possible function for this adaptation is that it helps the animal suck up water through the trunk.<ref name=snorkel /> Elephants inhale mostly through the trunk, although some air goes through the mouth. They have a [[Biology:Hindgut fermentation|hindgut fermentation]] system, and their large and small intestines together reach {{convert|35|m|ft|0|abbr=on}} in length. The majority of an elephant's food intake goes undigested despite the process lasting up to a day.<ref name=Shoshani78 />

{{anchor|Sex organs}}A male elephant's testes are located internally near the kidneys.<ref>{{cite journal | author1 = Short, R. V.| author2 = Mann, T.| author3 = Hay, Mary F. | year = 1967 | title = Male reproductive organs of the African elephant, Loxodonta africana | url = http://www.reproduction-online.org/content/13/3/517.full.pdf | journal = Journal of Reproduction and Fertility | volume = 13 | issue = 3| pages = 517–36 | doi = 10.1530/jrf.0.0130517 | pmid = 6029179 | doi-access = free }}</ref> The elephant's penis can reach a length of {{convert|100|cm|in|0|abbr=on}} and a diameter of {{convert|16|cm|in|0|abbr=on}} at the base. It is S-shaped when fully erect and has a Y-shaped orifice. The female has a well-developed clitoris at up to {{convert|40|cm|in|0|abbr=on}}. 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 [[Biology:Mammary gland|mammary gland]]s occupy the space between the front legs, which puts the suckling calf within reach of the female's trunk.<ref name=Shoshani78 /> Elephants have a unique organ, the [[Biology: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.<ref>Shoshani, p. 80.</ref> Females have also been observed with secretions from the temporal glands.<ref name=Sukumar149>Sukumar, p. 149.</ref>

===Body temperature ===
Elephants are [[Biology:Homeothermy|homeotherms]], and maintain their average body temperature at ~ 36&nbsp;°C, with minimum 35.2&nbsp;°C during cool season, and maximum 38.0&nbsp;°C during hot dry season.<ref name=":1">{{Cite journal|date=2018|title=Savanna elephants maintain homeothermy under African heat|url=https://link.springer.com/article/10.1007%2Fs00360-018-1170-5|journal=Journal of Comparative Physiology B|doi=10.1007/s00360-018-1170-5|last1=Mole|first1=Michael A.|last2=Rodrigues Dáraujo|first2=Shaun|last3=Van Aarde|first3=Rudi J.|last4=Mitchell|first4=Duncan|last5=Fuller|first5=Andrea|volume=188|issue=5|pages=889–897|pmid=30008137|s2cid=51626564}}</ref> Sweat glands are absent in the elephant's skin, but water diffuses through the skin, allowing cooling by evaporative loss.<ref>{{Cite journal|date=1984|title=Do elephants need to sweat?|journal=Journal of Zoology|doi=10.1080/02541858.1984.11447892|last1=Wright|first1=P.G.|last2=Luck|first2=C.P.|volume=19|issue=4|pages=270–274|doi-access=free}}</ref><ref>{{Cite journal|date=1970|title=The epidermis and its keratinisation in the African Elephant (Loxodonta Africana)|journal=Zoologica Africana|doi=10.1080/00445096.1970.11447400|doi-access=free|last1=Spearman|first1=R. I. C.|volume=5|issue=2|pages=327–338}}</ref><ref>{{Cite book|last=Sokolov|first=V.E.|title=Mammal Skin|publisher=University of California Press|year=1982|pages=449–453}}</ref> Other physiological or behavioral features may assist with thermoregulation such as flapping ears,<ref>{{Cite web|last=Hidden|first=PA|date=2009|title=Thermoregulation in African elephants (Loxodonta africana)|url=http://wiredspace.wits.ac.za/bitstream/handle/10539/7262/?sequence=1}}</ref> mud bathing, spraying water on the skin, seeking shade,<ref name=":1" /><ref>{{Cite journal|date=2012|title=Taking the heat: thermoregulation in Asian elephants under different climatic conditions|url=https://link.springer.com/article/10.1007/s00360-011-0609-8|journal=Journal of Comparative Physiology B Volume|doi=10.1007/s00360-011-0609-8|last1=Weissenböck|first1=Nicole M.|last2=Arnold|first2=Walter|last3=Ruf|first3=Thomas|volume=182|issue=2|pages=311–319|pmid=21938480|s2cid=16163113}}</ref> and adopting different walking patterns.<ref>{{Cite journal|date=2020|title=Temperature as a constraint on the timing and duration of African elephant foraging trips|url=https://academic.oup.com/jmammal/article-abstract/101/6/1670/5986389?redirectedFrom=fulltext|journal=Journal of Mammalogy|doi=10.1093/jmammal/gyaa129|last1=Rozen-Rechels|first1=David|last2=Valls-Fox|first2=Hugo|last3=Mabika|first3=Cheryl Tinashe|last4=Chamaillé-Jammes|first4=Simon|volume=101|issue=6|pages=1670–1679}}</ref> 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.<ref>{{Cite journal|date=2018|title=Locally-curved geometry generates bending cracks in the African elephant skin|journal=Nature Communications|doi=10.1038/s41467-018-06257-3|doi-access=free|last1=Martins|first1=António F.|last2=Bennett|first2=Nigel C.|last3=Clavel|first3=Sylvie|last4=Groenewald|first4=Herman|last5=Hensman|first5=Sean|last6=Hoby|first6=Stefan|last7=Joris|first7=Antoine|last8=Manger|first8=Paul R.|last9=Milinkovitch|first9=Michel C.|volume=9|issue=1|page=3865|pmid=30279508|pmc=6168576|bibcode=2018NatCo...9.3865M}}</ref>

== Behaviour and life history ==

=== Ecology and activities ===
{{multiple image
| align = right
| direction = vertical
| image1 = Elephant eating Yala Sri Lanka.ogv
| caption1 = An Asian elephant feeding on grass in Sri Lanka
| image2 = African elephant (Loxodonta africana) reaching up 1.jpg
| caption2 = An African bush elephant using its prehensile trunk for foraging
}}
The African bush elephant can be found in habitats as diverse as dry [[Earth:Savanna|savanna]]hs, [[Earth:Desert|desert]]s, [[Earth:Marsh|marsh]]es, and lake shores, and in elevations from sea level to mountains above the [[Earth:Snow line|snow line]]. Forest elephants mainly live in equatorial forests but will enter [[Biology:Gallery forest|gallery forest]]s and [[Earth:Ecotone|ecotone]]s between forests and savannahs.<ref name=Shoshani42>Shoshani, pp. 42–51.</ref> Asian elephants prefer areas with a mix of grasses, low woody plants, and trees, primarily inhabiting dry thorn-scrub forests in southern India and Sri Lanka and [[Earth:Evergreen forest|evergreen forest]]s in [[Biology:Malay Peninsula|Malaya]].<ref name=Asian>{{cite journal|author1=Shoshani, J. |author2=Eisenberg, J. F. |year=1982|title=''Elephas maximus''|journal=Mammalian Species|volume=182|issue=182|pages=1–8|url=http://www.science.smith.edu/msi/pdf/i0076-3519-182-01-0001.pdf|jstor=3504045|doi=10.2307/3504045}}</ref> Elephants are [[Earth:Herbivore|herbivorous]] and will eat leaves, twigs, fruit, bark, grass and roots.<ref name=Shoshani42 /> They are born with sterile intestines and require bacteria obtained from their mother's feces to digest vegetation.<ref>{{cite web|url=https://www.bbc.co.uk/nature/adaptations/Coprophagia |title= Dung eater |publisher=BBC Nature |access-date=27 November 2011}}</ref> African elephants are mostly browsers while Asian elephants are mainly [[Biology:Grazing|grazers]]. They can consume as much as {{convert|150|kg|lb|abbr=on}} of food and {{convert|40|L|gal|abbr=on}} of water in a day. Elephants tend to stay near water sources.<ref name=Shoshani42 /> 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.<ref name=Shoshani69 /><ref name=Shoshani124 /> Elephants average 3–4 hours of sleep per day.<ref name="Siegel">{{cite journal|author=Siegel, J.M.|s2cid=234089|year=2005|title=Clues to the functions of mammalian sleep|journal=Nature|volume=437|issue=7063|pages=1264–71|doi=10.1038/nature04285|pmid=16251951|bibcode=2005Natur.437.1264S}}</ref> Both males and family groups typically move {{convert|10|–|20|km|mi|0|abbr=on}} a day, but distances as far as {{convert|90|–|180|km|mi|0|abbr=on}} have been recorded in the Etosha region of Namibia. Elephants go on seasonal migrations in search of food, water, minerals, and mates.<ref>Sukumar, p. 159.</ref> At Chobe National Park, Botswana, herds travel {{convert|325|km|mi|0|abbr=on}} to visit the river when the local waterholes dry up.<ref>{{cite book|author=Hoare, B.|year=2009|title=Animal Migration: Remarkable Journeys in the Wild|publisher=University of California Press|pages=58–59|isbn=978-0-520-25823-5}}</ref>

Because of their large size, elephants have a huge impact on their environments and are considered [[Biology:Keystone species|keystone species]]. Their habit of uprooting trees and undergrowth can transform savannah into grasslands; when they dig for water during drought, they create waterholes that can be used by other animals. They can enlarge waterholes when they bathe and wallow in them. At [[Earth:Mount Elgon|Mount Elgon]], elephants excavate caves that are used by [[Biology:Ungulate|ungulate]]s, hyraxes, bats, birds and insects.<ref name=Shoshani226 /> Elephants are important [[Earth:Seed dispersal|seed dispersers]]; African forest elephants ingest and defecate seeds, with either no effect or a positive effect on [[Biology:Germination|germination]]. The seeds are typically dispersed in large amounts over great distances.<ref>{{cite journal|author1=Campos-Arceiz, A. |author2=Blake, S. |year=2011|title=Mega-gardeners of the forest – the role of elephants in seed dispersal|journal=Acta Oecologica|volume=37|issue=6|pages=542–53|doi=10.1016/j.actao.2011.01.014|url=http://faculty.washington.edu/timbillo/Readings%20and%20documents/CO2%20and%20Forests%20readings/Campos%20Arceiz%202011%20Elephant%20seed%20dispersal.pdf|bibcode=2011AcO....37..542C}}</ref> 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 [[Biology:Malayan tapir|tapir]].<ref>{{cite journal|author1=Campos-Arceiz, A. |author2=Traeholt, C. |author3=Jaffar, R. |author4=Santamaria, L. |author5=Corlett, R. T. |year=2012|title=Asian tapirs are no elephants when it comes to seed dispersal|journal=Biotropica|volume=44|issue=2|pages=220–27|doi=10.1111/j.1744-7429.2011.00784.x|hdl=10261/56573 }}</ref> Because most of the food elephants eat goes undigested, their dung can provide food for other animals, such as [[Biology:Dung beetle|dung beetle]]s and monkeys.<ref name=Shoshani226>Shoshani, pp. 226–29.</ref> 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 [[Earth:Surface runoff|run off]], leading to [[Earth:Erosion|erosion]].<ref name=Shoshani124>Eltringham, pp. 124–27.</ref>
[[File:Forest elephant.jpg|right|thumb|Forest elephant in habitat. It is considered to be an important seed disperser.]]
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.<ref name=Shoshani124 /> At Hluhluwe–Umfolozi Game Reserve, {{wipe|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.<ref>{{cite news|title=Elephants kill endangered rhino|work=BBC News|date=14 February 2000|url=http://news.bbc.co.uk/2/hi/642731.stm|access-date=5 October 2012}}</ref> The size of adult elephants makes them nearly invulnerable to predators.<ref name=Asian /> Calves may be preyed on by [[Biology:Lion|lion]]s, [[Biology:Spotted hyena|spotted hyena]]s, and wild dogs in Africa<ref name=African>{{cite journal|author1=Laursen, L.|author2=Bekoff, M.|year=1978|title=''Loxodonta africana''|journal=Mammalian Species|volume=92|pages=1–8|url=http://www.science.smith.edu/msi/pdf/i0076-3519-092-01-0001.pdf|jstor=3503889|doi=10.2307/3503889|issue=92|url-status=dead|archive-url=https://web.archive.org/web/20130430140501/http://www.science.smith.edu/msi/pdf/i0076-3519-092-01-0001.pdf|archive-date=30 April 2013|df=dmy-all}}</ref> and tigers in Asia.<ref name=Asian /> The lions of 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.<ref>{{cite journal |author1=Power, R. J. |author2=Shem Compion, R. X. |s2cid=86371484 |year=2009 |title=Lion predation on elephants in the Savuti, Chobe National Park, Botswana |journal=African Zoology |volume=44 |issue=1 |pages=36–44 |doi=10.3377/004.044.0104}}</ref><ref>{{cite journal |last=Joubert |first=D. |year=2006 |title=Hunting behaviour of lions (''Panthera leo'') on elephants (''Loxodonta africana'') in the Chobe National Park, Botswana |journal= African Journal of Ecology |volume=44 |issue=2 |pages=279–281 |doi=10.1111/j.1365-2028.2006.00626.x}}</ref> There are rare reports of adult Asian elephants falling prey to [[Biology:Tiger|tiger]]s.<ref>{{cite news |year=2009 |url=http://www.newindianexpress.com/cities/kochi/article103095.ece |title=Tiger kills elephant at Eravikulam park |newspaper=The New Indian Express}}</ref> Elephants appear to distinguish between the growls of larger predators like tigers and smaller predators like [[Biology:Leopard|leopard]]s (which have not been recorded killing calves); they react to leopards less fearfully and more aggressively.<ref>{{cite journal|author1=Thuppil, V.|author2=Coss, R. G.|year=2013|title=Wild Asian elephants distinguish aggressive tiger and leopard growls according to perceived danger|journal=Biology Letters|volume=9|issue=5|pages=20130518|doi=10.1098/rsbl.2013.0518|pmid=24026347|pmc=3971691|url=http://mon.univ-montp2.fr/claroline/backends/download.php?url=L1REXzE3LTEwLTIwMTMvVGh1cHBpbF9ldF9hbF8yMDEzLnBkZg%3D%3D&cidReset=true&cidReq=UMBGAE339|url-status=dead|archive-url=https://web.archive.org/web/20140714225616/http://mon.univ-montp2.fr/claroline/backends/download.php?url=L1REXzE3LTEwLTIwMTMvVGh1cHBpbF9ldF9hbF8yMDEzLnBkZg%3D%3D&cidReset=true&cidReq=UMBGAE339|archive-date=14 July 2014|df=dmy-all}}</ref> Elephants tend to have high numbers of parasites, particularly [[Biology:Nematode|nematode]]s, compared to other herbivores. This is due to lower predation pressures that would otherwise kill off many of the individuals with significant parasite loads.<ref>Sukumar, p. 121.</ref>

=== Social organisation ===
[[File:Serengeti Elefantenherde1.jpg|thumb|left|A family of African bush elephants]]
Female elephants spend their entire lives in tight-knit [[Social: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 [[Social:Matriarchy|matriarch]] which is often the eldest female.<ref name=Sukumar175 /> She remains leader of the group until death<ref name=African /> or if she no longer has the energy for the role;<ref name=Kingdon53>Kingdon, p. 53.</ref> a study on zoo elephants showed that when the matriarch died, the levels of faecal [[Chemistry:Corticosterone|corticosterone]] ('stress hormone') dramatically increased in the surviving elephants.<ref>{{cite web|author1=Harris, M. |author2=Sherwin, C. |author3=Harris, S. |url=http://randd.defra.gov.uk/Document.aspx?Document=WC05007_7719_FRP.pdf|title=Defra final report on elephant welfare|publisher=University of Bristol|year=2008}}</ref> When her tenure is over, the matriarch's eldest daughter takes her place; this occurs even if her sister is present.<ref name=African /> One study found that younger matriarchs are more likely than older ones to under-react to severe danger.<ref>{{cite journal|author1=McComb, K. |author2=Shannon, G. |author3=Durant, S. M. |author4=Sayialel, K. |author5=Slotow, R. |author6=Poole, J. |author7=Moss, C. |year=2011|title=Leadership in elephants: the adaptive value of age|journal=Proceedings of the Royal Society B: Biological Sciences|volume=278|issue=1722|pages=3270–76|doi=10.1098/rspb.2011.0168|url=http://www.lifesci.sussex.ac.uk/cmvcr/Publications_files/ElephantLeadersProofs.pdf |pmid=21411454 |pmc=3169024}}</ref> Family groups may split after becoming too large for the available resources.<ref>{{cite book|author1=Vaughan, T. |author2=Ryan, J. |author3=Czaplewski, N. |year=2011|title=Mammalogy|publisher=Jones & Bartlett Learning|page=136|isbn=978-0763762995}}</ref>

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.<ref name=Sukumar175>Sukumar, pp. 175–79.</ref>

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.<ref name=Sukumar175 />
[[File:Asian Elephant at Corbett National Park 15.jpg|thumb|right|Lone bull: Adult male elephants spend much of their time alone or in single-sex groups]]
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.<ref name=Sukumar179 />

Male elephants can be quite sociable when not competing for dominance or mates, and will form long-term relationships.<ref name="O'Connell-Rodwell">{{cite web|author=O'Connell-Rodwell, C.|date=November 2010|title=How Male Elephants Bond|publisher=Smithsonian.com|access-date=4 June 2017|url=http://www.smithsonianmag.com/science-nature/how-male-elephants-bond-64316480/}}</ref> A [[Dominance hierarchy|dominance hierarchy]] exists among males, whether they range socially or solitarily. Dominance depends on the age, size and sexual condition,<ref name=Sukumar179 /> 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,<ref name="O'Connell-Rodwell"/> whose presence appears to control their aggression and prevent them from exhibiting "deviant" behaviour.<ref>{{cite journal|author1=Slotow, R. |author2=van Dyk, G. |author3=Poole, J. |author4=Page, B. |author5=Klocke, A. |s2cid=136330 |year=2000|title=Older bull elephants control young males|journal=Nature|volume=408|issue=6811|pages=425–26|doi=10.1038/35044191|pmid=11100713|bibcode=2000Natur.408..425S}}</ref> Adult males and females come together for reproduction. Bulls associate with family groups if an oestrous cow is present.<ref name=Sukumar179>Sukumar, pp. 179–83.</ref>

=== Sexual behaviour ===

==== Musth ====
[[File:2005-tusker-musth-crop.jpg|thumb|right|upright|Bull in musth]]

Adult males enter a state of increased [[Chemistry:Testosterone|testosterone]] known as musth. In a population in southern India, males first enter musth at the age of 15, but it is not very intense until they are older than 25. At Amboseli, bulls under 24 do not go into musth, while half of those aged 25–35 and all those over 35 do. Young bulls appear to enter musth during the dry season (January–May), while older bulls go through it during the wet season (June–December). The main characteristic of a bull's musth is a fluid secreted from the [[Biology:Temporin|temporal gland]] that runs down the side of his face. He may urinate with his penis still in his sheath, which causes the urine to spray on his hind legs. Behaviours associated with musth include walking with the head held high and swinging, picking at the ground with the tusks, marking, rumbling and waving only one ear at a time. This can last from a day to four months.<ref name=Sukumar100 />

Males become extremely aggressive during musth. Size is the determining factor in [[Biology: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.<ref name=Sukumar100>Sukumar, pp. 100–08.</ref>

==== Mating ====
[[File:Elephantsmating.jpg|thumb|Bull mating with a member of a female group]]

Elephants are [[Social:Polygyny|polygynous]] breeders,<ref>Sukumar, p. 89.</ref> and [[Biology:Animal sexual behaviour|copulations]] are most frequent during the peak of the wet season.<ref>Sukumar, p. 262.</ref> A cow in oestrus releases chemical signals ([[Biology:Pheromone|pheromone]]s) 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 [[Biology:Flehmen response|flehmen response]], which requires the male to collect a chemical sample with his trunk and bring it to the [[Biology:Vomeronasal organ|vomeronasal organ]].<ref>Sukumar, pp. 98–99.</ref> The oestrous cycle of a cow lasts 14–16 weeks with a 4–6-week [[Biology:Follicular phase|follicular phase]] and an 8- to 10-week luteal phase. While most mammals have one surge of [[Biology:Luteinizing hormone|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.<ref>{{cite web|title=Elephant Reproduction Project: The Estrous Cycle of Elephants|publisher=Smithsonian National Zoo|access-date=8 October 2012|url=http://nationalzoo.si.edu/scbi/ReproductiveScience/ElephantBreedRepro/EstrousCycle.cfm|url-status=dead|archive-url=https://web.archive.org/web/20120606110832/http://nationalzoo.si.edu/SCBI/ReproductiveScience/ElephantBreedRepro/EstrousCycle.cfm|archive-date=6 June 2012|df=dmy-all}}</ref> Fertility rates in cows decline around 45–50 years of age.<ref name=Kingdon53 />

Bulls engage in a behaviour known as mate-guarding, where they follow oestrous females and defend them from other males.<ref>{{cite journal | author = Poole Joyce H | s2cid = 53150105 | year = 1989 | title = Mate guarding, reproductive success and female choice in African elephants | url = https://www.researchgate.net/publication/222028670 | format = PDF | journal = Animal Behaviour | volume = 37 | pages = 842–849 | doi=10.1016/0003-3472(89)90068-7}}</ref> Most mate-guarding is done by musth males, and females actively seek to be guarded by them, particularly older ones.<ref>Sukumar, p. 113.</ref> Thus these bulls have more reproductive success.<ref name=Sukumar179 /> Musth appears to signal to females the condition of the male, as weak or injured males do not have normal musths.<ref>Sukumar, p. 117.</ref> For young females, the approach of an older bull can be intimidating, so her relatives stay nearby to provide support and reassurance.<ref name=Shoshani106 /> During copulation, the male lays his trunk over the female's back.<ref>Kingdon, p. 69.</ref> The penis is very mobile, being able to move independently of the pelvis.<ref name="FowlerMikota2006">{{cite book|author1=Murray E. Fowler|author2=Susan K. Mikota|title=Biology, Medicine, and Surgery of Elephants|url=https://books.google.com/books?id=oCpiZA61tyQC&q=elephant+penis+mating&pg=PA353|year=2006|publisher=John Wiley & Sons|isbn=978-0-8138-0676-1|page=353}}</ref> Prior to mounting, it curves forward and upward. Copulation lasts about 45 seconds and does not involve pelvic thrusting or ejaculatory pause.<ref name=Estes>{{cite book|author=Estes, R.|title=The behavior guide to African mammals: including hoofed mammals, carnivores, primates|url=https://archive.org/details/isbn_0520080858|url-access=registration|pages=[https://archive.org/details/isbn_0520080858/page/263 263–66]|year=1991|publisher=University of California Press|isbn=978-0-520-08085-0}}</ref> Elephant sperm must swim close to {{convert|2|m|ft|abbr=on}} to reach the egg. By comparison, human sperm has to swim around only {{convert|76.2|mm|in|abbr=on}}.<ref>{{cite book|author=Sims, M.|year=2009|title=In the Womb: Animals|publisher=National Geographic Books|page=118|isbn=978-1426201752}}</ref>

[[Biology: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 [[Biology:Animal sexual behaviour#Autoeroticism or masturbation|masturbate one another]] with their trunks.<ref>{{cite book|author=Bagemihl, B.|year=1999|title=Biological Exuberance: Animal Homosexuality and Natural Diversity|url=https://archive.org/details/biologicalexuber00bage|url-access=registration|publisher=St. Martin's Press|pages=[https://archive.org/details/biologicalexuber00bage/page/427 427–30]|isbn=978-1-4668-0927-7}}</ref>

=== Birth and development ===
[[File:Loxodontacyclotis.jpg|thumb|An African forest elephant mother bathing with her calf]]
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.<ref>Sukumar, pp. 259–62.</ref> Calves are born {{convert|85|cm|in|abbr=on}} tall and weigh around {{convert|120|kg|lb|abbr=on}}.<ref name=Shoshani106>Moss, pp. 106–13.</ref> Typically, only a single young is born, but twins sometimes occur.<ref name=pregnancy /><ref>{{cite web|url=http://news.am/eng/videos/825.html|title=Rare elephant twins born in northern Kwa-Zulu Natal|work=news.am}}</ref> The relatively long pregnancy is maintained by five [[Biology:Corpus luteum|corpus luteum]]s (as opposed to one in most mammals) and gives the foetus more time to develop, particularly the brain and trunk.<ref name="pregnancy">{{cite journal|author1=Lueders, I.|author2=Niemuller, C. |author3=Rich, P. |author4=Gray, C. |author5=Hermes, R. |author6=Goeritz, F. |author7=Hildebrandt, T. B. |year=2012|title=Gestating for 22 months: luteal development and pregnancy maintenance in elephants|journal=Proceedings of the Royal Society B: Biological Sciences|volume=279|issue=1743|pages=3687–96|pmid=22719030|doi=10.1098/rspb.2012.1038|pmc= 3415912}}</ref> As such, newborn elephants are precocial and quickly stand and walk to follow their mother and family herd.<ref name=Sukumar126 /> 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.<ref name=Shoshani106 />

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.<ref name=Sukumar126 />

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.<ref name=Sukumar126>Sukumar, pp. 126–29.</ref>

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<ref name=Shoshani106 /> while the latter become mature around 14–15 years.<ref name=Sukumar179 /> Adulthood starts at about 18 years of age in both sexes.<ref>{{cite web|title=Elephant Life Cycle|publisher=ElephantsForever.co.za|url=http://www.elephantsforever.co.za/life-cycle.html |access-date=21 June 2014}}</ref><ref>{{cite web|title=Elephant Life Cycle – Adolescence|publisher=ElephantsForever.co.za|url=http://www.elephantsforever.co.za/life-cycle-adolescence.html |access-date=21 June 2014}}</ref> Elephants have long lifespans, reaching 60–70 years of age.<ref name=Shoshani70 /> Lin Wang, a captive male Asian elephant, lived for 86 years.<ref>{{cite news|title=War veteran elephant dies|work=BBC News|date=26 February 2003|url=http://news.bbc.co.uk/2/hi/asia-pacific/2800737.stm|access-date=8 January 2013}}</ref>

=== Communication ===
{{main article|Biology:Elephant communication}}
[[File:Three elephant's curly kisses.jpg|thumb|Asian elephants greeting each other by inter-twining their trunks]]
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.<ref>Payne and Langbauer, p. 116.</ref>

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.<ref>Payne and Langbauer, pp. 119–20.</ref>
[[File:Loxodonta africana oral rumble visualized with acoustic camera (25fps) - pone.0048907.s003.ogv|thumb|right|Low frequency rumble visualised with acoustic camera]]
Elephants produce several vocalisations, usually through the larynx, though some may be modified by the trunk.<ref name=Shoshani120>Payne and Langbauer, pp. 120–21.</ref> These include trumpets, roars, barks, snorts, growls and rumbles which may be produced for either short or long range communication.<ref>Sukumar, p. 141.</ref> Elephants may produce [[Physics:Infrasound|infrasonic]] rumbles.<ref name="infrasonic">{{cite journal| author1= Herbest, C. T.| author2 = Stoeger, A.| author3 = Frey, R.| author4 = Lohscheller, J.| author5 = Titze, I. R.| author6 = Gumpenberger, M.| author7 = Fitch, W. T.|s2cid=32792564|year=2012|title=How Low Can You Go? Physical Production Mechanism of Elephant Infrasonic Vocalizations|journal= Science|volume=337|issue=6094|pages=595–599|doi=10.1126/science.1219712|pmid=22859490|bibcode=2012Sci...337..595H}}</ref> For Asian elephants, these calls have a frequency of 14–24&nbsp;[[Hertz|Hz]], with [[Physics:Sound pressure|sound pressure]] levels of 85–90&nbsp;[[Physics:Decibel|dB]] and last 10–15 seconds.<ref name="Payne">{{cite journal|author1=Payne, K.B. |author2=Langbauer, W.R. |author3=Thomas, E.M. |s2cid=1480496 |year=1986|title=Infrasonic calls of the Asian elephant (''Elephas maximus'')|journal=Behavioral Ecology and Sociobiology|volume=18|issue=4|pages=297–301|doi=10.1007/BF00300007}}</ref> For African elephants, calls range from 15 to 35&nbsp;Hz with sound pressure levels as high as 117&nbsp;dB, allowing communication for many kilometres, with a possible maximum range of around {{convert|10|km|0|abbr=on}}.<ref name="Larom">{{cite journal|author1=Larom, D.|author2=Garstang, M. |author3=Payne, K. |author4=Raspet, R. |author5=Lindeque, M. |year=1997|title=The influence of surface atmospheric conditions on the range and area reached by animal vocalizations|journal=Journal of Experimental Biology|volume=200|pages=421–31|pmid=9057305|url=http://jeb.biologists.org/cgi/reprint/200/3/421.pdf|issue=Pt 3|doi=10.1242/jeb.200.3.421 }}</ref>

Elephants are known to [[Physics: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 {{convert|32|km|mi|abbr=on}}. Seismic waveforms produced from predator alarm calls travel {{convert|16|km|mi|0|abbr=on}}.<ref>{{cite journal|author1=O'Connell-Rodwell, C. E.|author2=Wood, J. D. |author3=Rodwell, T. C. |author4=Puria, S. |author5=Partan, S. R. |author6=Keefe, R. |author7=Shriver, D. |author8=Arnason, B. T. |author9=Hart, L. A. |s2cid=33221888 |year=2006 |title=Wild elephant (''Loxodonta africana'') breeding herds respond to artificially transmitted seismic stimuli |journal=Behavioural and Ecological Sociobiology |volume=59 |pages=842–50 |doi=10.1007/s00265-005-0136-2 |url=http://www.sscnet.ucla.edu/anthro/bec/papers/OConnellRodwell_4.16.07.pdf |issue=6 |url-status=dead |archive-url=https://web.archive.org/web/20131203071054/http://www.sscnet.ucla.edu/anthro/bec/papers/OConnellRodwell_4.16.07.pdf |archive-date=3 December 2013 }}</ref><ref>{{cite journal|author1=O'Connell-Rodwell C. E. |author2=Arnason, B. |author3=Hart, L. A. |year=2000|title=Seismic properties of Asian elephant (''Elephas maximus'') vocalizations and locomotion|journal=Journal of the Acoustical Society of America|volume=108|issue=6|pages=3066–72|pmid=11144599|doi=10.1121/1.1323460|bibcode=2000ASAJ..108.3066O}}</ref>

=== Intelligence and cognition ===
<!-- Please do not add any more examples to this section. This subject already has its own article -->
[[File:Insightful-Problem-Solving-in-an-Asian-Elephant-pone.0023251.s005.ogv|thumb|upright=1.15|right|Elephant rolling a block to allow it to reach food]]

Elephants exhibit [[Philosophy:Mirror test|mirror self-recognition]], an indication of [[Philosophy:Self-awareness|self-awareness]] and [[Cognition|cognition]] that has also been demonstrated in some [[Biology:Ape|ape]]s and [[Biology:Dolphin|dolphin]]s.<ref>{{cite journal|author1=Plotnik, J. M. |author2=de Waal, F. B. M. |author3=Reiss, D. |name-list-style=amp |year=2006|title=Self-recognition in an Asian elephant|journal=Proceedings of the National Academy of Sciences|volume=103|issue=45|pages=17053–57|doi=10.1073/pnas.0608062103|pmid=17075063 |bibcode = 2006PNAS..10317053P |pmc=1636577|doi-access=free }}</ref> 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.<ref>{{cite journal|author=Rensch, B.|year=1957|title=The intelligence of elephants|journal=Scientific American|volume=196|issue=2|pages=44–49|doi=10.1038/scientificamerican0257-44|bibcode=1957SciAm.196b..44R}}</ref> Elephants are among the [[Biology:Tool use by animals|species known to use tools]]. An Asian elephant has been observed modifying branches and using them as [[Engineering:Fly-killing device|flyswatters]].<ref>{{cite journal|author1=Hart, B. J. |author2=Hart, L. A. |author3=McCory, M. |author4=Sarath, C. R. |s2cid=53184282 |year=2001|title=Cognitive behaviour in Asian elephants: use and modification of branches for fly switching|journal=Animal Behaviour|volume=62|issue=5|pages=839–47|doi=10.1006/anbe.2001.1815}}</ref> Tool modification by these animals is not as advanced as that of [[Biology:Pan (genus)|chimpanzee]]s. Elephants are popularly thought of as having an excellent memory. This could have a factual basis; they possibly have [[Cognitive map|cognitive map]]s 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.<ref name="cognition">{{cite journal|author1=Byrne, R. W.|author2=Bates, L.|author3=Moss C. J.|year=2009|title=Elephant cognition in primate perspective|journal= Comparative Cognition & Behavior Reviews|volume=4|pages=65–79|doi=10.3819/ccbr.2009.40009|df=dmy-all|doi-access=free}}</ref>

Scientists debate the extent to which elephants feel [[Philosophy:Emotion|emotion]]. They appear to show interest in the bones of their own kind, regardless of whether they are related.<ref>{{cite journal|author1=McComb, K. |author2=Baker, L. |author3=Moss, C. |year=2006|title=African elephants show high levels of interest in the skulls and ivory of their own species|journal=Biology Letters|volume=2|issue=1|pages=26–28|doi=10.1098/rsbl.2005.0400|pmid=17148317|pmc=1617198}}</ref> As with chimps and dolphins, a dying or dead elephant may elicit attention and aid from others, including those from other groups. This has been interpreted as expressing "concern";<ref>{{cite journal|author1=Douglas-Hamilton, I.|author2=Bhallaa, S.|author3=Wittemyera, G.|author4=Vollratha, F.|year=2006|title=Behavioural reactions of elephants towards a dying and deceased matriarch|journal=Applied Animal Behaviour Science|volume=100|issue=1|pages=87–102|doi=10.1016/j.applanim.2006.04.014|url=http://www.savetheelephants.org/files/pdf/publications/2006%20Douglas-Hamilton%20et%20al%20Behavioural%20reactions%20of%20elephants%20to%20a%20dying%20matriarch.pdf|url-status=dead|archive-url=https://web.archive.org/web/20120511045326/http://www.savetheelephants.org/files/pdf/publications/2006%20Douglas-Hamilton%20et%20al%20Behavioural%20reactions%20of%20elephants%20to%20a%20dying%20matriarch.pdf|archive-date=11 May 2012|df=dmy-all}}</ref> however, others would dispute such an interpretation as being anthropomorphic;<ref>{{cite web|url=https://www.sciencedaily.com/releases/2009/06/090611065839.htm|title=What really prompts the dog's 'Guilty Look'|website=Science Daily|year=2009|access-date=5 September 2013}}</ref><ref>{{cite web|url=http://www.psychologytoday.com/blog/animal-emotions/200906/anthropomorphic-double-talk-can-animals-be-happy-not-unhappy-no|title=Anthropomorphic Double-Talk: Can Animals Be Happy But Not Unhappy? No!|author=Bekoff, M.|year=2009|access-date=5 September 2013}}</ref> 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".<ref>{{cite book |title= When Elephants Weep: Emotional Lives of Animals|last= Masson|first= Jeffrey Moussaieff|author2=Susan McCarthy |year= 1996|publisher= Vintage|isbn= 978-0-09-947891-1 |page= 272}}</ref>

== Conservation ==

=== Status ===
{{multiple image|caption_align=center|header_align=center
| direction = horizontal
| header = Distribution of elephants
| align = top
| width1 = 205
| image1 = African Elephant distribution map.svg
| caption1 = African bush and forest elephants
| width2 = 183
| image2 = Asian Elephant area.png
| caption2 = Asian elephant
}}
African bush elephants were listed as Endangered by the [[Organization:International Union for Conservation of Nature|International Union for Conservation of Nature]] (IUCN) in 2021,<ref name="IUCN">{{cite iucn |title=''Loxodonta africana.'' |author1=Gobush, K.S. |author2=Edwards, C.T.T. |author3=Maisels, F. |author4=Wittemeyer, G. |author5=Balfour, D. |author6=Taylor, R.D. |date=2021 |page=e.T181008073A181022663 |access-date=25 March 2021 |url=https://www.iucnredlist.org/species/181008073/181022663}}</ref> and African forest elephants were listed as [[Biology:Critically Endangered|Critically Endangered]] in the same year.<ref>{{cite iucn |title=''Loxodonta cyclotis'' |author1=Gobush, K.S. |author2=Edwards, C.T.T. |author3=Maisels, F. |author4=Wittemeyer, G. |author5=Balfour, D. |author6=Taylor, R.D. |name-list-style=amp |date=2021 |page=e.T181007989A181019888 |access-date=25 March 2021}}</ref> In 1979, Africa had an estimated minimum population of 1.3&nbsp;million elephants, with a possible upper limit of 3.0&nbsp;million. By 1989, the population was estimated to be 609,000; with 277,000 in Central Africa, 110,000 in [[Place:East Africa|eastern Africa]], 204,000 in [[Place:Southern Africa|southern Africa]], and 19,000 in [[Place: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.<ref>Douglas-Hamilton, pp. 178–82.</ref> Conversely, studies in 2005 and 2007 found populations in eastern and southern Africa were increasing by an average annual rate of 4.0%.<ref name=IUCN /> The IUCN estimated that total population in Africa is estimated at around to 415,000 individuals for both species combined as of 2016.<ref>{{cite web|author=Thouless, C.R.; Dublin, H.T.; Blanc, J.J.; Skinner, D.P.; Daniel, T.E.; Taylor, R.D.; Maisels, F.; Frederick, H. L.; Bouché, P.|year=2016|title=African Elephant Status Report 2016|publisher=IUCN African Elephant Database|accessdate=2021-06-05|url=https://portals.iucn.org/library/sites/library/files/documents/SSC-OP-060_A.pdf}}</ref>

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 [[Biology: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 [[Biology: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, 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.<ref name=IUCN /> 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.<ref name="StarNews">Star website. [http://www.the-star.co.ke/news/2016/06/04/first-lady-launches-campaign-to-save-elephants_c1363294 "First Lady launches campaign to save elephants."] 4 June 2016. Accessed 5 June 2016.</ref>

In 2020, the IUCN listed the Asian elephant as endangered due to an almost 50% population decline over "the last three generations"<ref name="IUCN2">{{cite iucn |author1=Williams, C. |display-authors=0 |year=2020 |url=https://www.iucnredlist.org/species/7140/45818198 |title=''Elephas maximus'' |access-date=2020-06-03}}</ref> Asian elephants once ranged from [[Place:Syria|Syria]] and Iraq (the subspecies [[Biology:Syrian elephant|''Elephas maximus asurus'']]), to China (up to the Yellow River)<ref name=Shoshani174>Daniel, p. 174.</ref> and Java. It is now extinct in these areas,<ref name=IUCN2 /> and the current range of Asian elephants is highly fragmented.<ref name=Shoshani174 /> 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.<ref name=IUCN2 />

=== Threats ===
[[File:African elephant carcass ratio, OWID.svg|thumb|upright=1.3|African elephant carcass ratio as of 2015<ref>{{cite web |title=African elephant carcass ratio |url=https://ourworldindata.org/grapher/african-elephant-carcass-ratio |website=Our World in Data |access-date=15 February 2020}}</ref>]]
[[File:Ivory trade.jpg|thumb|upright|Men with elephant tusks at [[Place:Dar es Salaam|Dar es Salaam]], Tanzania, c. 1900]]

The [[Biology:Poaching|poaching]] of elephants for their ivory, meat and hides has been one of the major threats to their existence.<ref name=IUCN2 /> Historically, numerous cultures made ornaments and other works of art from elephant ivory, and its use rivalled that of gold.<ref name=Shoshani202>Martin, pp. 202–07</ref> The ivory trade contributed to the African elephant population decline in the late 20th century.<ref name=IUCN /> 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.<ref name=Shoshani202 /> Around the same time, Kenya destroyed all its ivory stocks.<ref name="ivory">{{cite magazine|author=Christy, B.|date=October 2012|title=Ivory Worship|magazine=National Geographic|url=http://ngm.nationalgeographic.com/2012/10/ivory/christy-text|access-date=17 October 2012}}</ref> 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.<ref name=Shoshani202 /> 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.<ref name=ivory />

The ban allowed the elephant to recover in parts of Africa.<ref name=Shoshani202 /> In January 2012, 650 elephants in Bouba Njida National Park, Cameroon, were killed by Chadian raiders.<ref>{{Cite news|last1=Hicks|first1=Celeste|title=86 elephants killed in Chad poaching massacre|url=https://www.theguardian.com/environment/2013/mar/19/86-elephants-chad-poaching-massacre|newspaper=The Guardian|access-date=23 September 2015|date=19 March 2013}}</ref> This has been called "one of the worst concentrated killings" since the ivory ban.<ref name="ivory" /> 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.<ref name=IUCN2 /> 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.<ref>{{Cite news|author=Ryan, F.|url=https://www.theguardian.com/environment/2015/sep/26/china-and-us-agree-on-ivory-ban-in-bid-to-end-illegal-trade-globally|title=China and US agree on ivory ban in bid to end illegal trade globally|date=26 September 2015|access-date=12 October 2015|newspaper=The Guardian}}</ref>

Other threats to elephants include [[Earth:Habitat destruction|habitat destruction]] and [[Biology: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 [[Biology:Wildlife corridor|wildlife corridor]]s which gave the animals greater space and maintain the long term viability of large populations.<ref name=IUCN2 />

== Association with humans ==

=== Working animal ===
[[File:Elefant pune.jpg|thumb|Working elephant as transport]]

Elephants have been [[Biology:Working animal|working animal]]s since at least the Indus Valley Civilization<ref>Sukumar, p. 57.</ref> 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.<ref name=Shoshani149 /> They were traditionally captured with traps and lassos, but since 1950, [[Chemistry:Sedative|tranquillisers]] have been used.<ref>Wylie, pp. 120–23.</ref>

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.<ref name="Shoshani149" /> In northern Thailand, the animals are used to digest coffee beans for Black Ivory coffee.<ref>{{Cite news|author=Topper, R|date=15 October 2012|title=Elephant Dung Coffee: World's Most Expensive Brew Is Made With Pooped-Out Beans|work=The Huffington Post|access-date=10 December 2012|url=http://www.huffingtonpost.com/2012/10/15/elephant-dung-coffee-black-ivory_n_1968096.html}}</ref> 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.<ref name="Shoshani149">McNeely, pp. 149–50.</ref> Musth bulls can be difficult and dangerous to work with and are chained and semi-starved until the condition passes.<ref>Easa, p. 86.</ref> In India, many working elephants are alleged to have been subject to abuse. They and other captive elephants are thus protected under The Prevention of Cruelty to Animals Act of 1960.<ref>{{cite web|author1=Bist, S. S. |author2=Cheeran, J. V. |author3=Choudhury, S. |author4=Barua, P. |author5=Misra, M. K. |title=The domesticated Asian elephant in India|publisher= Regional Office for Asia and the Pacific|access-date=25 December 2012|url=http://www.fao.org/docrep/005/ad031e/ad031e0g.htm}}</ref>

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.<ref>{{cite news |author=Thomas Fuller |title=Unemployed, Myanmar's Elephants Grow Antsy, and Heavier |url=https://www.nytimes.com/2016/01/31/world/asia/myanmar-logging-elephants-unemployment.html |work={{wipe|The New York Times}} |date=30 January 2016 |access-date=31 January 2016 }}</ref><ref>{{cite web |url=http://www.fao.org/docrep/005/ad031e/ad031e0r.htm |title=The elephant situation in Thailand and a plea for co-operation |author=Roger Lohanan |date= February 2001 |access-date=31 January 2016 |publisher=FAO }}</ref>

The practice of working elephants has also been attempted in Africa. The taming of African elephants in the Belgian Congo began by decree of Leopold II of Belgium during the 19th century and continues to the present with the Api Elephant Domestication Centre.<ref>Smith, pp. 152–54.</ref>

=== Warfare ===
<!-- Please do not add any more examples to this section. This subject already has its own article -->
[[File:Schlacht bei Zama Gemälde H P Motte.jpg|thumb|Battle of Zama by Henri-Paul Motte, 1890]]

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.<ref name=Shoshani146 />

One of the earliest references to war elephants is in the Indian epic ''[[Religion:Mahabharata|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 [[History:Nanda Empire|Nandas]] (5th and 4th centuries BC) army while 9,000 may have been used in the [[History:Maurya Empire|Mauryan army]] (between the 4th and 2nd centuries BC). The ''[[Religion:Arthashastra|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.<ref>Sukumar, pp. 59–64.</ref> From South Asia, the use of elephants in warfare spread west to [[Religion:Iran|Persia]]<ref name=Shoshani146 /> and east to Southeast Asia.<ref name=SEA /> The Persians used them during the [[Place:Achaemenid Empire|Achaemenid Empire]] (between the 6th and 4th centuries BC)<ref name=Shoshani146 /> while Southeast Asian states first used war elephants possibly as early as the 5th century BC and continued to the 20th century.<ref name="SEA">{{cite book|author=Griffin, B|year=2004|contribution=Elephants: From the Sacred to the Mundane|title=Southeast Asia: A Historical Encyclopedia, from Angkor Wat to East Timor|volume=1 |editor=Gin Ooi, K.|pages=487–89|isbn=978-1-57607-770-2}}</ref>

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.<ref name=":0">{{Cite book|title=Warfare in Ancient Greece|last=Everson|first=Tim|publisher=The History Press|year=2004|isbn=978-0-7524-9506-4|location=Brimscombe Port}}</ref> Alexander trained his foot soldiers to injure the animals and cause them to panic during wars with both the Persians and Indians. Ptolemy, who was one of Alexander's generals, used corps of Asian elephants during his reign as the ruler of Egypt (which began in 323 BC). His son and successor Ptolemy II (who began his rule in 285 BC) obtained his supply of elephants further south in [[Place:Nubia|Nubia]]. From then on, war elephants were employed in the Mediterranean and North Africa throughout the [[History:Classical antiquity|classical period]]. The Greek king Pyrrhus used elephants in his attempted invasion of Rome in 280 BC. While they frightened the Roman horses, they were not decisive and Pyrrhus ultimately lost the battle. The Carthaginian general Hannibal took elephants across the [[Place:Alps|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.<ref name="Shoshani146">Wylie (2000), pp. 146–48.</ref>

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 [[History:Parthian Empire|Parthians]].<ref name=":0" />

=== Zoos and circuses ===
[[File:Flickr - …trialsanderrors - Terrific flights over ponderous elephants, poster for Forepaugh ^ Sells Brothers, ca. 1899.jpg|thumb|right|Circus poster, c. 1900]]
Elephants were historically kept for display in the menageries of Ancient Egypt, [[Biology:Elephants in ancient China|China]], {{wipe|Ancient Greece}}, and [[History:Ancient Rome|Rome]]. The Romans in particular pitted them against humans and other animals in gladiator events. In the [[History:Modern history|modern era]], elephants have traditionally been a major part of [[Biology:Zoo|zoo]]s and circuses around the world. In circuses, they are trained to perform tricks. The most famous circus elephant was probably Jumbo (1861 – 15 September 1885), who was a major attraction in the Barnum & Bailey Circus.<ref>Shoshani, pp. 168–69.</ref> 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.<ref name=Shoshani184>Tuttle, pp. 184–88.</ref>

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.<ref>{{cite web|author=Sterm, A.|date=28 February 2005|title=Elephant deaths at zoos reignite animal debate: Zoo supporters cite conservation, activists cite confined spaces|publisher=MSNBC/Reuters|url=http://www.elephants.com/media/MSNBC_2_28_05.htm|access-date=24 October 2012}}</ref> Elephants have been recorded displaying [[Biology: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.<ref>{{cite web|author1=Harris, M. |author2=Sherwin, C. |author3=Harris, S. |url=http://randd.defra.gov.uk/Document.aspx?Document=WC05007_7719_FRP.pdf|title=Defra Final Report on Elephant Welfare|publisher=University of Bristol|date=10 November 2008|access-date=16 November 2011}}</ref> 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.<ref>{{cite web|author=Mott, M.|date=11 December 2008|title=Wild elephants live longer than their zoo counterparts|publisher=National Geographic News|url=http://news.nationalgeographic.com/news/2008/12/081211-zoo-elephants.html|access-date=24 October 2012}}</ref>

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.<ref>{{cite web|title=Circus Myths: The true cruelty under the big top|publisher=Humane Society of the United States|url=http://www.humanesociety.org/issues/circuses_entertainment/facts/circus_myths.html|date=25 September 2009|access-date=24 October 2012}}</ref> In testimony to a US federal court in 2009, Barnum & Bailey Circus CEO Kenneth Feld acknowledged that circus elephants are struck behind their ears, under their chins and on their legs with metal-tipped prods, called bull hooks or ankus. Feld stated that these practices are necessary to protect circus workers and acknowledged that an elephant trainer was reprimanded for using an electric shock device, known as a hot shot or electric prod, on an elephant. Despite this, he denied that any of these practices harm elephants.<ref>{{cite news|author=Pickler, N.|url=http://www.dailyrecord.com/article/20090304/UPDATES01/903040429/Circus-CEO-says-elephants-struck-not-hurt|title=Circus CEO says elephants are struck, but not hurt|agency=Associated Press|date=4 March 2009|access-date=25 October 2012|url-status=dead|archive-url=https://archive.today/20130120082119/http://www.dailyrecord.com/article/20090304/UPDATES01/903040429/Circus-CEO-says-elephants-struck-not-hurt|archive-date=20 January 2013|df=dmy-all}}</ref> 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.<ref>Wylie, p. 142.</ref> Ringling Bros. and Barnum and Bailey circus retired its touring elephants in May 2016.<ref>{{cite news|last1=Karimi|first1=Faith|title=Ringling Bros. elephants perform last show |url=http://www.cnn.com/2016/05/01/us/ringling-bros-elephants-last-show/index.html|access-date=21 September 2017|publisher=CNN|date=2 May 2016|location=St. Petersburg, Florida}}</ref>

=== Attacks ===
Elephants can exhibit bouts of aggressive behaviour and engage in destructive actions against humans.<ref name="huggler">{{cite news | url = https://www.independent.co.uk/environment/animal-behaviour-rogue-elephants-419678.html | archive-url = https://web.archive.org/web/20080514154342/http://www.independent.co.uk/environment/animal-behaviour-rogue-elephants-419678.html | url-status = dead | archive-date = 14 May 2008 | title = Animal Behaviour: Rogue Elephants | author = Huggler, J. | work= The Independent | date = 12 October 2006 | access-date = 16 June 2007 | location=London}}</ref> 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.<ref name="teenage">{{cite news | url = http://www.smh.com.au/articles/2006/02/16/1140064206413.html | title = Elephant rage: they never forgive, either | author = Highfield, R. | work = The Sydney Morning Herald| date = 17 February 2006 | access-date = 16 June 2007}}</ref><ref>{{cite news|author=Siebert, C.|date=8 October 2006|title=An Elephant Crackup?|work=Nytimes.com|url=https://www.nytimes.com/2006/10/08/magazine/08elephant.html|access-date=25 October 2012}}</ref> 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.<ref name="huggler" />
Local people have reported their belief that some elephants were drunk during their attacks, although officials have disputed this explanation.<ref>{{cite news | url = http://news.bbc.co.uk/1/hi/world/south_asia/241781.stm | title = India elephant rampage | work = BBC News | date = 24 December 1998 | access-date = 16 June 2007}}</ref><ref>{{cite news | url = http://news.bbc.co.uk/1/hi/world/south_asia/482001.stm | title = Drunken elephants trample village | work = BBC News | date = 21 October 1999 | access-date = 16 June 2007}}</ref> 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.<ref>{{cite news | url = http://news.bbc.co.uk/2/hi/south_asia/2583891.stm | title = Drunk elephants kill six people | work = BBC News | date = 17 December 2002 | access-date = 16 June 2007}}</ref>

=== Cultural depictions ===
<!-- Please do not add any more examples to this section. This subject already has its own article -->
[[File:Blind monks examining an elephant.jpg|thumb|left|Parable of the elephant and the blind monks; illustrated by Hanabusa Itchō. (Ukiyo-e woodcut, 1888)]]
In many cultures, elephants represent strength, power, wisdom, longevity, stamina, leadership, sociability, nurturance and loyalty.<ref>{{cite book|last=Guading|first=Madonna|title=The signs and symbols bible : the definitive guide to mysterious markings|year=2009|publisher=Sterling Pub. Co|location=New York|isbn=978-1402770043|pages=239|url=https://books.google.com/books?id=SImTll3uupIC&pg=PA239}}</ref><ref>{{cite book|title=Nature's Ways Lore, Legend, Fact and Fiction.|year=2006|publisher=F+W Media|page=37|isbn=9780715333938}}</ref><ref>{{cite web|title=Elephant: The Animal and Its Ivory in African Culture |url=http://www.fowler.ucla.edu/category/exhibitions-education/elephant-animal-and-its-ivory-african-culture |publisher=Fowler Museum at UCLA |access-date=24 January 2013 |url-status=dead |archive-url=https://web.archive.org/web/20130330072035/http://www.fowler.ucla.edu/category/exhibitions-education/elephant-animal-and-its-ivory-african-culture |archive-date=30 March 2013 }}</ref> 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.<ref name="Van Riper 73" /> The expression "[[Social:Elephant in the room|elephant in the room]]" refers to an obvious truth that is ignored or otherwise unaddressed.<ref>{{cite book|title=Cambridge Academic Content Dictionary Paperback with CD-ROM|publisher=Cambridge University Press|page=[https://archive.org/details/cambridgeacademi00camb/page/298 298]|isbn=978-0-521-69196-3|date=3 November 2008|url=https://archive.org/details/cambridgeacademi00camb/page/298}}</ref> The story of the [[Blind men and an elephant|blind men and an elephant]] teaches that reality can be observed from different perspectives.<ref>{{cite book|author=Nevid, J. S.|year=2008|title=Psychology: Concepts and Applications|publisher=Wadsworth Publishing|page=477|isbn=978-0-547-14814-4}}</ref>

Elephants have been represented in art since [[Earth:Paleolithic|Paleolithic]] times. Africa, in particular, contains many rock paintings and engravings of the animals, especially in the Sahara and southern Africa.<ref>Wylie, pp. 63–65.</ref> In Asia, the animals are depicted as motifs in [[Religion:Hinduism|Hindu]] and [[Religion:Buddhism|Buddhist]] shrines and temples.<ref name=Shoshani158 /> Elephants were often difficult to portray by people with no first-hand experience of them.<ref>Kingdon, p. 31.</ref> The [[History:Ancient Rome|ancient Romans]], who kept the animals in captivity, depicted anatomically accurate elephants on [[Engineering:Mosaic|mosaic]]s in Tunisia and Sicily. At the beginning of the [[History:Middle Ages|Middle Ages]], when Europeans had little to no access to the animals, elephants were portrayed more like fantasy creatures. They were often depicted with horse- or bovine-like bodies with trumpet-like trunks and tusks like a boar; some were even given hooves. Elephants were commonly featured in motifs by the stonemasons of the Gothic churches. As more elephants began to be sent to European kings as gifts during the 15th century, depictions of them became more accurate, including one made by [[Biography:Leonardo da Vinci|Leonardo da Vinci]]. Despite this, some Europeans continued to portray them in a more stylised fashion.<ref>Wylie, pp. 83–84.</ref> Max Ernst's 1921 surrealist painting, ''The Elephant Celebes,'' depicts an elephant as a [[Engineering:Silo|silo]] with a trunk-like hose protruding from it.<ref>{{cite book|author1=Klinsöhr-Leroy, C. |author2=Grosenick, U. |year=2004|title=Surrealism|publisher=Taschen|page=50|isbn=978-3-8228-2215-9}}</ref>
[[File:Illustration at p. 73 in Just So Stories (c1912).png|thumb|right|Woodcut illustration for "The Elephant's Child" by Rudyard Kipling]]
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.<ref name=Shoshani158>McNeely, pp. 158–60.</ref> Similar ideas existed among other African societies, who believed that their chiefs would be [[Philosophy:Reincarnation|reincarnated]] as elephants. During the 10th century AD, the people of Igbo-Ukwu, near the Niger Delta, buried their leaders with elephant tusks.<ref>Wylie, p. 79.</ref> The animals' religious importance is only totemic in Africa<ref>Sukumar, p. 87.</ref> 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.<ref name=Shoshani158 /> One of the most important Hindu deities, the elephant-headed [[Biology:Ganesha|Ganesha]], is ranked equal with the supreme gods [[Religion:Shiva|Shiva]], [[Religion:Vishnu|Vishnu]], and [[Religion:Brahma|Brahma]].<ref>Sukumar, p. 64.</ref> Ganesha is associated with writers and merchants and it is believed that he can give people success as well as grant them their desires.<ref name=Shoshani158 /> In Buddhism, [[Biography:Gautama Buddha|Buddha]] is said to have been a white elephant reincarnated as a human.<ref>Sukumar, p. 62.</ref> In [[Religion:Islam|Islam]]ic tradition, the year 570 when [[Biography:Muhammad|Muhammad]] was born is known as the [[Religion:Year of the Elephant|Year of the Elephant]].<ref>{{cite book|author=Haykal, M. H.|year=2008|title=The Life of Muḥammad|publisher=Islamic Book Trust|page=52|isbn=978-983-9154-17-7}}</ref> Elephants were thought to be religious themselves by the Romans, who believed that they worshipped the sun and stars.<ref name=Shoshani158 />

Elephants are ubiquitous in Western [[Social:Popular culture|popular culture]] as emblems of the exotic, especially since – as with the [[Biology:Giraffe|giraffe]], [[Biology:Hippopotamus|hippopotamus]] and rhinoceros – there are no similar animals familiar to Western audiences.<ref name="Van Riper 73" /> The use of the elephant as a symbol of the U.S. Republican Party began with an 1874 cartoon by Thomas Nast.<ref>{{cite web|url=http://www.harpweek.com/09Cartoon/BrowseByDateCartoon.asp?Year=2003&Month=November&Date=7 |title=Cartoon of the Day: "The Third-Term Panic" |publisher=HarpWeek |access-date=1 September 2008 |url-status=dead |archive-url=https://web.archive.org/web/20110921045800/http://www.harpweek.com/09Cartoon/BrowseByDateCartoon.asp?Year=2003&Month=November&Date=7 |archive-date=21 September 2011 |df=dmy }}</ref> 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'', [[Company:The Walt Disney Company|Disney]]'s ''Dumbo,'' and Kathryn and Byron Jackson's ''The Saggy Baggy Elephant''. Other elephant heroes given human qualities include Jean de Brunhoff's Babar, David McKee's Elmer, and Dr. Seuss's Horton.<ref name="Van Riper 73">{{cite book|author1=Van Riper |author2=A. B. |title=Science in Popular Culture: A Reference Guide|publisher=Greenwood Press|year=2002|pages=73–75|isbn=978-0-313-31822-1}}</ref>

== See also ==
* [[Biology:Animal track|Animal track]]
* Beehive fences use elephants' fear of bees to minimise conflict with humans
* Desert elephant
* [[Unsolved:Elephants' graveyard|Elephants' graveyard]]
* [[Biology:Motty|Motty]], captive hybrid of an Asian and African elephant
* National Elephant Day (Thailand)

== References ==
{{Reflist|30em}}

=== Bibliography ===
{{Refbegin|60em}}
* {{cite book |editor=Shoshani, J. |year=2000 |title=Elephants: Majestic Creatures of the Wild |publisher=Checkmark Books |isbn=978-0-87596-143-9 |oclc=475147472}}
** {{cite book |author1=Shoshani, J. |author2=Shoshani, S. L. |chapter=What Is an Elephant? |title=Elephants: Majestic Creatures of the Wild |pages=14–15}}
** {{cite book |author=Shoshani, J. |chapter=Comparing the Living Elephants |title=Elephants: Majestic Creatures of the Wild |pages=36–51}}
** {{cite book |author=Shoshani, J. |chapter=Anatomy and Physiology |title=Elephants: Majestic Creatures of the Wild |pages=66–80}}
** {{cite book |author=Easa, P. S. |chapter=Musth in Asian Elephants |title=Elephants: Majestic Creatures of the Wild |pages=85–86}}
** {{cite book |author=Moss, C |chapter=Elephant Calves: The Story of Two Sexes |title=Elephants: Majestic Creatures of the Wild |pages=106–13}}
** {{cite book |author1=Payne, K. B. |author2=Langauer, W. B |chapter=Elephant Communication |title=Elephants: Majestic Creatures of the Wild |pages=116–23}}
** {{cite book |author=Eltringham, S. K. |chapter=Ecology and Behavior |title=Elephants: Majestic Creatures of the Wild |pages=124–27}}
** {{cite book |author=Wylie, K. C. |chapter=Elephants as War Machines |title=Elephants: Majestic Creatures of the Wild |pages=146–48}}
** {{cite book |author=McNeely, J. A. |chapter=Elephants as Beasts of Burden |title=Elephants: Majestic Creatures of the Wild |pages=149–50}}
** {{cite book |author=Smith, K. H. |chapter=The Elephant Domestication Centre of Africa |title=Elephants: Majestic Creatures of the Wild |pages=152–54}}
** {{cite book |author=McNeely, J. A. |chapter=Elephants in Folklore, Religion and Art |title=Elephants: Majestic Creatures of the Wild |pages=158–65}}
** {{cite book |author=Shoshani, S. L. |chapter=Famous Elephants |title=Elephants: Majestic Creatures of the Wild |pages=168–71}}
** {{cite book |author=Daniel, J. C. |chapter=The Asian Elephant Population Today |title=Elephants: Majestic Creatures of the Wild |pages=174–77}}
** {{cite book |author=Douglas-Hamilton, I. |chapter=The African Elephant Population Today |title=Elephants: Majestic Creatures of the Wild |pages=178–83}}
** {{cite book |author=Tuttle, C. D. |chapter=Elephants in Captivity |title=Elephants: Majestic Creatures of the Wild |pages=184–93}}
** {{cite book |author=Martin, E. B. |chapter=The Rise and Fall of the Ivory Market |title=Elephants: Majestic Creatures of the Wild |pages=202–07}}
** {{cite book |author=Shoshani, J. |chapter=Why Save Elephants? |title=Elephants: Majestic Creatures of the Wild |pages=226–29}}
* {{cite book|author= Sukumar, R.|title= The Living Elephants: Evolutionary Ecology, Behaviour, and Conservation|url= https://archive.org/details/livingelephantse00suku_0|url-access= registration|date= 11 September 2003|publisher= Oxford University Press, USA|isbn= 978-0-19-510778-4 |oclc= 935260783}}
* {{cite book|author =Kingdon, J.|title= East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals|url= https://books.google.com/books?id=xdH_Gi92-fAC |date= 29 December 1988|publisher= University of Chicago Press|isbn= 978-0-226-43722-4 |oclc= 468569394}}
* {{cite book|author= Wylie, D.|title= Elephant|url= https://books.google.com/books?id=Aj75snWdfv8C |date= 15 January 2009|publisher= Reaktion Books|isbn= 978-1-86189-615-5 |oclc= 740873839}}
{{Refend}}

== Further reading ==
* {{cite book |last=Carrington |first=Richard |title=Elephants: A Short Account of their Natural History, Evolution and Influence on Mankind| publisher=Chatto & Windus|year=1958|oclc=911782153}}
* Nance, Susan (2013). ''Entertaining Elephants: Animal Agency and the Business of the American Circus''. Baltimore, MD: Johns Hopkins University Press, 2013.
* Saxe, John Godfrey (1872). "The Blindmen and the Elephant" at [[Wikisource]].  ''The Poems of John Godfrey Saxe''.
* {{cite book | last=Williams | first=Heathcote | title=Sacred Elephant | location=New York | publisher=Harmony Books | year=1989 | isbn=978-0-517-57320-4}}

== External links ==
<!-- ==============================({{NoMoreLinks}})============================== -->
<!-- DO ''not'' ADD MORE LINKS TO THIS ARTICLE. WIKIPEDIA IS ''not'' A COLLECTION OF LINKS -->
<!-- If you think that your link might be useful, instead of placing it here, put -->
<!-- it on this article's discussion page first. Links that have not been verified -->
<!--                                WILL BE DELETED                                -->
<!-- ============================================================================= -->
* [https://elephantconservation.org International Elephant Foundation]


[[Category:Paraphyletic groups]]

{{Sourceattribution|Elephant|1}}