View source for Elephant

{{Short description|Largest living land animal}}
{{Hatnote|This article is primarily about living elephants, for their close extinct relatives, see [[Elephantidae]], for other uses, see [[Elephant (disambiguation)]]}}
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{{Use dmy dates|date=January 2020}}
{{Use British English|date=January 2013}}
{{Paraphyletic group
| name = Elephants
| image = {{CSS image crop
   |Image = African Bush Elephant.jpg
   |bSize = 240
   |cWidth = 220
   |cHeight = 290
   |oTop = 65
   |oLeft = 10
   }}
| image_caption = A female [[African bush elephant]] in [[Mikumi National Park]], Tanzania
| auto = yes
| fossil_range = {{Fossil range|Late Miocene | Present}}
| parent = Elephantidae
| includes = * ''[[African elephant|Loxodonta]]'' <small>Anonymous, 1827</small>
* ''[[Elephas]]'' <small>[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]</small>
* {{extinct}}''[[Palaeoloxodon]]'' <small>Matsumoto, 1925</small>
| excludes = * {{extinct}}''[[Mammuthus]]'' <small>Brookes, 1828</small>
| range_map = Loxodonta Elephas distribution.png
| range_map_caption = Distribution of living elephant species
}}

'''Elephants''' are the [[Largest and heaviest animals|largest]] living land animals. Three living [[species]] are currently recognised: the [[African bush elephant]] (''[[Loxodonta]] africana''), the [[African forest elephant]] (''L. cyclotis''), and the [[Asian elephant]] (''[[Elephas]] maximus''). They are the only surviving members of the [[Family (biology)|family]] [[Elephantidae]] and the [[Order (biology)|order]] [[Proboscidea]]; extinct relatives include [[mammoth]]s and [[mastodon]]s. Distinctive features of elephants include a long [[proboscis]] called a trunk, [[tusk]]s, large ear flaps, pillar-like legs, and tough but sensitive grey skin. The trunk is [[Prehensility|prehensile]], bringing food and water to the mouth and grasping objects. Tusks, which are derived from the incisor teeth, serve both as weapons and as tools for moving objects and digging. The large ear flaps assist in maintaining a constant body temperature as well as in communication. African elephants have larger ears and concave backs, whereas Asian elephants have smaller ears and convex or level backs.

Elephants are scattered throughout [[sub-Saharan Africa]], South Asia, and Southeast Asia and are found in different habitats, including [[savanna]]hs, forests, deserts, and [[marsh]]es. They are [[herbivorous]], and they stay near water when it is accessible. They are considered to be [[keystone species]], due to their impact on their environments. Elephants have a [[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 leader of a female group, usually the oldest cow, is known as the [[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 [[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 [[Elephant communication|communicate]] by touch, sight, smell, and sound; elephants use [[infrasound]] and [[seismic communication]] over long distances. [[Elephant cognition|Elephant intelligence]] has been compared with that of [[primate]]s and [[cetacea]]ns. They appear to have [[self-awareness]], and possibly show concern for dying and dead individuals of their kind.

African bush elephants and Asian elephants are listed as [[Endangered species (IUCN status)|endangered]] and African forest elephants as [[critically endangered]] on the [[IUCN Red List]]s. One of the biggest threats to elephant populations is the [[ivory trade]], as the animals are [[Poaching|poached]] for their ivory tusks. Other threats to wild elephants include [[habitat destruction]] and conflicts with local people. Elephants are used as [[working animal]]s in Asia. In the past, they were used in war; today, they are often controversially put on display in zoos, or employed for entertainment in [[circus]]es. Elephants have an iconic status [[Cultural depictions of elephants|in human culture]] and have been widely featured in art, folklore, religion, literature, and popular culture.

==Etymology==
The word ''elephant'' is derived from the [[Latin]] word {{lang|la|elephas}} ([[genitive]] {{lang|la|elephantis}}) {{gloss|elephant}}, which is the [[Latinisation of names|Latinised]] form of the [[ancient Greek]] {{lang|grc|ἐλέφας}} ({{transliteration|grc|elephas}}) (genitive {{lang|grc|ἐλέφαντος}} ({{transliteration|grc|elephantos}},<ref name=LSJ>{{cite book |author=Liddell, H. G. |author2=Scott, R |year=1940 |title=A Greek-English Lexicon |edition=Revised and augmented |place=Oxford |publisher=Clarendon Press |chapter=ἐλέφας |chapter-url=https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0057:entry=e)le/fas}}</ref>)) probably from a non-[[Indo-European language]], likely [[Phoenician language|Phoenician]].<ref name=etymology>{{cite encyclopedia|author=Harper, D. |title=Elephant |dictionary=Online Etymology Dictionary|url=http://www.etymonline.com/index.php?term=elephant|access-date=25 October 2012 |archive-date=24 December 2013|archive-url=https://web.archive.org/web/20131224141504/http://www.etymonline.com/index.php?term=elephant |url-status=live}}</ref> It is attested in [[Mycenaean Greek]] as {{transliteration|gmy|e-re-pa}} (genitive {{transliteration|gmy|e-re-pa-to}}) in [[Linear B]] syllabic script.<ref>{{Cite book |editor1=Nosch, M.-L. |editor2=Laffineur, R. |title=Kosmos. Jewellery, Adornment and Textiles in the Aegean Bronze Age |publisher=Peeters |series=Aegaeum 33 |place=Liège |year=2012 |last1=Lujan |first1=E. R. |last2=Bernabe |first2=A. |chapter=Ivory and horn production in Mycenaean texts |chapter-url=https://www.academia.edu/2229199 |access-date=22 January 2013 |archive-date=20 October 2021 |archive-url=https://web.archive.org/web/20211020172849/https://www.academia.edu/2229199 |url-status=live}}</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 |archive-date=4 December 2012|archive-url=https://web.archive.org/web/20121204075314/http://www.palaeolexicon.com/default.aspx?static=12&wid=189|url-status=live}}</ref> As in Mycenaean Greek, [[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 [[Middle English]] as {{lang|enm|olyfaunt}} in {{Circa|1300}} and was borrowed from [[Old French]] {{lang|fro|oliphant}} in the 12th century.<ref name=etymology/>

==Taxonomy==
{{cladogram|style=font-size:70%|align=right|caption=A cladogram of the elephants within [[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|access-date=19 June 2017|archive-date=24 February 2021|archive-url=https://web.archive.org/web/20210224122358/http://phylodiversity.net/azanne/csfar/images/d/d9/Afrotherian_mammals.pdf|url-status=usurped | issn=0025-1461}}</ref>
|{{clade
|label1=[[Afrotheria]]
|1={{Clade
   |label1=[[Afroinsectiphilia]]
   |1={{Clade
      |label1=[[Tubulidentata]]
                 |1=[[Orycteropodidae]] [[File:Aardvark2 (PSF) colourised.png|60 px]]
      |label2=[[Afroinsectivora]]
      |2={{Clade
         |label1=[[Macroscelidea]]
                    |1=[[Macroscelididae]] [[File:Rhynchocyon chrysopygus-J Smit white background.jpg|50 px]]
         |label2=[[Afrosoricida]]
                     |2={{Clade
                         |1=[[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=[[Tenrecidae]] [[File:Brehms Thierleben - Allgemeine Kunde des Thierreichs (1876) (Tenrec ecaudatus).jpg|50 px]]
      }} }} }}
   |label2=[[Paenungulata]]
   |2={{Clade
      |label1=[[Hyrax|Hyracoidea]]
                 |1=[[Hyrax|Procaviidae]] [[File:DendrohyraxEminiSmit white background.jpg|50 px]]
      |label2=[[Tethytheria]]
      |2={{Clade
         |label1=[[Proboscidea]]
                    |1=[[Elephantidae]] [[File:Elephas africanus - 1700-1880 - Print - Iconographia Zoologica - (white background).jpg|70 px]]
         |label2=[[Sirenia]]
                     |2={{Clade
                         |1=[[Dugongidae]] [[File:Dugong dugon Hardwicke white background.jpg|60 px]]
                         |2=[[Trichechidae]] [[File:Manatee white background.jpg|60 px]]
         }} }} }} }} }}}}
Elephants belong to the family [[Elephantidae]], the sole remaining family within the order [[Proboscidea]]. Their closest [[extant taxon|extant]] relatives are the [[sirenia]]ns ([[dugong]]s and [[manatee]]s) and the [[hyrax]]es, with which they share the [[clade]] [[Paenungulata]] within the superorder [[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|issue=1 |page=6|doi=10.1186/1471-2148-7-6 |pmid=17244368 |pmc=1784077 |doi-access=free |bibcode=2007BMCEE...7....6K }}</ref> Elephants and sirenians are further grouped in the clade [[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 living elephants are recognised; the [[African bush elephant]] (''[[Loxodonta]] africana''), [[African forest elephant|forest elephant]] (''Loxodonta cyclotis''), and [[Asian elephant]] (''[[Elephas]] maximus'').<ref name=MSW3>{{cite book|author=Shoshani, J.|year=2005|contribution=Order Proboscidea|editor1=Wilson, D. E.|editor2=Reeder, D. M|url=https://books.google.com/books?id=JgAMbNSt8ikC&pg=PA91|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|access-date=11 November 2016|archive-date=1 February 2015|archive-url=https://web.archive.org/web/20150201164505/http://www.google.com/books?id=JgAMbNSt8ikC&pg=PA91|url-status=live}}</ref> [[African elephant]]s 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 |doi-access=free }}</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> [[Mammoth]]s (''Mammuthus'') are nested within living elephants as they are more closely related to Asian elephants than to African elephants.<ref>{{Cite journal |last1=Roca |first1=Alfred L. |last2=Ishida |first2=Yasuko |last3=Brandt |first3=Adam L. |last4=Benjamin |first4=Neal R. |last5=Zhao |first5=Kai |last6=Georgiadis |first6=Nicholas J. |date=2015 |title=Elephant Natural History: A Genomic Perspective |journal=Annual Review of Animal Biosciences |volume=3 |issue=1 |pages=139–167 |doi=10.1146/annurev-animal-022114-110838 |pmid=25493538}}</ref> Another extinct genus of elephant, ''[[Palaeoloxodon]]'', is also recognised, which appears to have close affinities with African elephants and to have hybridised with African forest elephants.<ref name="Palkopoulou">{{Cite journal |last1=Palkopoulou |first1=Eleftheria |last2=Lipson |first2=Mark |last3=Mallick |first3=Swapan |last4=Nielsen |first4=Svend |last5=Rohland |first5=Nadin |last6=Baleka |first6=Sina |last7=Karpinski |first7=Emil |last8=Ivancevic |first8=Atma M. |last9=To |first9=Thu-Hien |last10=Kortschak |first10=R. Daniel |last11=Raison |first11=Joy M. |date=2018-03-13 |title=A comprehensive genomic history of extinct and living elephants |journal=Proceedings of the National Academy of Sciences |language=en |volume=115 |issue=11 |pages=E2566–E2574 |doi=10.1073/pnas.1720554115 |issn=0027-8424 |pmc=5856550 |pmid=29483247 |bibcode=2018PNAS..115E2566P |doi-access=free}}</ref> Some species of the extinct ''[[Palaeoloxodon]]'' were even larger, all exceeding 4 metres in height and 10 tonnes in body mass, with ''[[Palaeoloxodon namadicus|P. namadicus]]'' being a contender for the largest land mammal to have ever existed.<ref name="Larramendi, A. 2015" />

===Evolution===
The earliest members of Proboscidea like ''[[Eritherium]]'' are known from the [[Paleocene]] of Africa, around 60 million years ago, the earliest proboscideans were much smaller than living elephants, with ''Eritherium'' having a body mass of around {{cvt|3-8|kg}}.<ref name="Gheerbrant2">{{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 |bibcode=2009PNAS..10610717G |doi=10.1073/pnas.0900251106 |pmc=2705600 |pmid=19549873 |doi-access=free}}</ref> By the late Eocene, some members of Proboscidea like ''[[Barytherium]]'' had reached considerable size, with an estimated mass of around 2 tonnes,<ref name="Larramendi, A. 2015" /> while others like ''[[Moeritherium]]'' are suggested to have been semi-aquatic.<ref>{{Cite journal |last1=Liu |first1=Alexander G. S. C. |last2=Seiffert |first2=Erik R. |last3=Simons |first3=Elwyn L. |date=2008-04-15 |title=Stable isotope evidence for an amphibious phase in early proboscidean evolution |journal=Proceedings of the National Academy of Sciences |language=en |volume=105 |issue=15 |pages=5786–5791 |bibcode=2008PNAS..105.5786L |doi=10.1073/pnas.0800884105 |issn=0027-8424 |pmc=2311368 |pmid=18413605 |doi-access=free}}</ref>

{{cladogram|style=font-size:70%|caption=Proboscidea phylogeny based on morphological and DNA evidence<ref>{{cite journal|last1=Baleka|first1=S. |last2=Varela |first2=L. |last3=Tambusso|first3=P. S.|last4=Paijmans|first4=J. L. A.|last5=Mothé|first5=D. |last6=Stafford Jr.|first6=T. W. |last7=Fariña|first7=R. A. |last8=Hofreiter |first8=M. |year=2022|title=Revisiting proboscidean phylogeny and evolution through total evidence and palaeogenetic analyses including ''Notiomastodon'' ancient DNA |journal=iScience |volume=25|issue=1 |page=103559 |doi=10.1016/j.isci.2021.103559 |pmid=34988402 |pmc=8693454 |bibcode=2022iSci...25j3559B}}</ref><ref>{{Cite book |last1=Benoit |first1=J. |chapter=Paleoneurology of the Proboscidea (Mammalia, Afrotheria): Insights from Their Brain Endocast and Labyrinth |date=2023 |title=Paleoneurology of Amniotes |pages=579–644 |editor-last=Dozo |editor-first=M. T. |place=Cham |publisher=Springer International Publishing |doi=10.1007/978-3-031-13983-3_15 |isbn=978-3-031-13982-6 |last2=Lyras |first2=G. A. |last3=Schmitt |first3=A. |last4=Nxumalo |first4=M. |last5=Tabuce |first5=R. |last6=Obada |first6=T. |last7=Mararsecul |first7=V. |last8=Manger |first8=P. |editor2-last=Paulina-Carabajal |editor2-first=A. |editor3-last=Macrini |editor3-first=T. E. |editor4-last=Walsh |editor4-first=S.}}</ref><ref name=Palkopoulou/>
|align=right
|{{clade
|label1=[[Proboscidea]]
|1={{clade
|1=early proboscideans, e.g. ''[[Moeritherium]]'' [[File:Moeritherium NT crop.jpg|70px]]
|2={{clade
   |1=[[Deinotheriidae]] [[File:Deinotherium12.jpg|70px]]
   |label2=[[Elephantiformes]]
   |2={{clade
      |1=[[Mammutidae]] <span style="{{MirrorH}}">[[File:BlankMastodon.jpg|70px]]</span>
       |2={{clade
         |1=[[Gomphotheriidae]] <span style="{{MirrorH}}">[[File:Gomphotherium NT small.jpg|70px]]</span>
         |2={{clade
            |1=[[Stegodontidae]] [[File:Stegodon ganesaDB.jpg|85px]]
            |label2=[[Elephantidae]]
            |2={{clade
            |1={{clade
               |1=''[[Loxodonta]]'' <span style="{{MirrorH}}">[[File:African elephant (PSF).png|60px]]</span>
               |2=''[[Palaeoloxodon]]'' [[File:Palaeoloxodon namadicus-bpk.jpg|60px]]
}}
               |2={{clade
                  |1=''[[Mammuthus]]'' [[File:Mammuthus trogontherii122DB.jpg|70px]]
                  |2=''[[Elephas]]'' <span style="{{MirrorH}}">[[File:Indian elephant (PSF).png|70px]]</span>
                  }}
               }}
            }}
         }}
      }}
   }}
 }}
}}
}}
A major event in proboscidean evolution was the collision of Afro-Arabia with Eurasia, during the Early Miocene, around 18–19 million years ago, allowing proboscideans to disperse from their African homeland across Eurasia and later, around 16–15 million years ago into North America across the [[Beringia|Bering Land Bridge]]. Proboscidean groups prominent during the Miocene include the [[deinothere]]s, along with the more advanced [[Elephantimorpha|elephantimorphs]], including [[Mammutidae|mammutids]] (mastodons), [[gomphothere]]s, [[Amebelodontidae|amebelodontids]] (which includes the "shovel tuskers" like ''[[Platybelodon]]''), [[Choerolophodontidae|choerolophodontids]] and [[Stegodontidae|stegodontids]].<ref name=Cantalapiedra-2021>{{Cite journal |last1=Cantalapiedra |first1=J. L. |last2=Sanisidro |first2=Ó. |last3=Zhang |first3=H. |last4=Alberdi |first4=M. T. |last5=Prado |first5=J. L. |last6=Blanco |first6=F. |last7=Saarinen |first7=J. |date=2021 |title=The rise and fall of proboscidean ecological diversity |url=https://www.nature.com/articles/s41559-021-01498-w |journal=Nature Ecology & Evolution |volume=5 |issue=9 |pages=1266–1272 |doi=10.1038/s41559-021-01498-w |pmid=34211141 |bibcode=2021NatEE...5.1266C |s2cid=235712060|hdl=10261/249360 |hdl-access=free }}</ref> Around 10 million years ago, the earliest members of the family [[Elephantidae]] emerged in Africa, having originated from gomphotheres.<ref>{{cite book |author1=Saegusa, H. |author2=Nakaya, H. |name-list-style=amp |author3=Kunimatsu, Y. |author4=Nakatsukasa, M. |author5=Tsujikawa, H. |author6=Sawada, Y. |author7=Saneyoshi, M. |author8=Sakai, T. |year=2014 |chapter=Earliest elephantid remains from the late Miocene locality, Nakali, Kenya |page=175 |chapter-url=https://apo.ansto.gov.au/dspace/bitstream/10238/9340/2/icmr_volume_low.pdf#page=188 |editor1=Kostopoulos, D. S. |editor2=Vlachos, E.  |editor3=Tsoukala, E. |title=VIth International Conference on Mammoths and Their Relatives |volume=102 |location=Thessaloniki |publisher=School of Geology, Aristotle University of Thessaloniki |isbn=978-960-9502-14-6}}</ref>

Elephantids are distinguished from earlier proboscideans by a major shift in the molar morphology to parallel lophs rather than the cusps of earlier proboscideans, allowing them to become higher-crowned (hypsodont) and more efficient in consuming grass.<ref>{{Cite journal |last=Lister |first=A. M. |date=2013 |title=The role of behaviour in adaptive morphological evolution of African proboscideans |doi=10.1038/nature12275 |journal=Nature |volume=500 |issue=7462 |pages=331–334 |pmid=23803767 |bibcode=2013Natur.500..331L |s2cid=883007}}</ref> The Late Miocene saw major climactic changes, which resulted in the decline and extinction of many proboscidean groups.<ref name="Cantalapiedra-2021" /> The earliest members of the modern genera of elephants (''Elephas'', ''Loxodonta'') as well as mammoths, appeared in Africa during the latest Miocene–early Pliocene around 7-4 million years ago.<ref>{{Cite book |last=Sanders |first=William J. |url=https://www.taylorfrancis.com/books/9781315118918 |title=Evolution and Fossil Record of African Proboscidea |date=2023-07-07 |publisher=CRC Press |isbn=978-1-315-11891-8 |edition=1 |location=Boca Raton |pages=154, 220 |language=en |doi=10.1201/b20016}}</ref> The elephantid genera ''Elephas'' (which includes the living Asian elephant) and ''Mammuthus'' (mammoths) migrated out of Africa during the late Pliocene, around 3.6 to 3.2 million years ago.<ref>{{Cite journal |last1=Iannucci |first1=Alessio |last2=Sardella |first2=Raffaele |date=2023-02-28 |title=What Does the "Elephant-''Equus''" Event Mean Today? Reflections on Mammal Dispersal Events around the Pliocene-Pleistocene Boundary and the Flexible Ambiguity of Biochronology |journal=Quaternary |volume=6 |issue=1 |page=16 |doi=10.3390/quat6010016 |doi-access=free |hdl=11573/1680082 |hdl-access=free}}</ref>

Over the course of the [[Early Pleistocene]], all non-elephantid probobscidean genera outside of the Americas became extinct with the exception of ''[[Stegodon]]'',<ref name="Cantalapiedra-2021" /> with gomphotheres dispersing into South America as part of the [[Great American interchange]],<ref name="Mothé et al 2016 (In Press)">{{cite journal |last1=Mothé |first1=Dimila |last2=dos Santos Avilla |first2=Leonardo |last3=Asevedo |first3=Lidiane |last4=Borges-Silva |first4=Leon |last5=Rosas |first5=Mariane |last6=Labarca-Encina |first6=Rafael |last7=Souberlich |first7=Ricardo |last8=Soibelzon |first8=Esteban |last9=Roman-Carrion |first9=José Luis |last10=Ríos |first10=Sergio D. |last11=Rincon |first11=Ascanio D. |last12=Cardoso de Oliveira |first12=Gina |last13=Pereira Lopes |first13=Renato |date=30 September 2016 |title=Sixty years after 'The mastodonts of Brazil': The state of the art of South American proboscideans (Proboscidea, Gomphotheriidae) |url=http://bibdigital.epn.edu.ec/bitstream/15000/17075/1/Moth%c3%a9%20et%20al.%2c%202016%20-%20Sixty%20years%20proboscideans.pdf |journal=Quaternary International |volume=443 |pages=52–64 |bibcode=2017QuInt.443...52M |doi=10.1016/j.quaint.2016.08.028}}</ref> and mammoths migrating into North America around 1.5 million years ago.<ref>{{Cite journal |last1=Lister |first1=A. M. |last2=Sher |first2=A. V. |date=2015 |title=Evolution and dispersal of mammoths across the Northern Hemisphere |journal=Science |volume=350 |issue=6262 |pages=805–809 |bibcode=2015Sci...350..805L |doi=10.1126/science.aac5660 |pmid=26564853 |s2cid=206639522|doi-access=free }}</ref> At the end of the Early Pleistocene, around 800,000 years ago the elephantid genus ''[[Palaeoloxodon]]'' dispersed outside of Africa, becoming widely distributed in Eurasia.<ref>{{Cite book |last=Lister |first=A. M. |chapter=Ecological Interactions of Elephantids in Pleistocene Eurasia |date=2004 |chapter-url=https://www.researchgate.net/publication/264788794 |title=Human Paleoecology in the Levantine Corridor |pages=53–60 |publisher=Oxbow Books |isbn=978-1-78570-965-4}}</ref> Proboscideans were represented by around 23 species at the beginning of the [[Late Pleistocene]]. Proboscideans underwent a dramatic decline during the Late Pleistocene as part of the [[Late Pleistocene extinctions]] of most large mammals globally, with all remaining non-elephantid proboscideans (including ''Stegodon'', [[mastodon]]s, and the American gomphotheres ''[[Cuvieronius]]'' and ''[[Notiomastodon]]'') and ''[[Palaeoloxodon]]'' becoming extinct, with mammoths only surviving in [[Relict (biology)|relict]] populations on islands around the [[Bering Strait]] into the Holocene, with their latest survival being on [[Wrangel Island]], where they persisted until around 4,000 years ago.<ref name=Cantalapiedra-2021/><ref>{{Cite journal |last1=Rogers |first1=R. L. |last2=Slatkin |first2=M. |date=2017 |title=Excess of genomic defects in a woolly mammoth on Wrangel island |journal=PLOS Genetics |volume=13 |issue=3 |page=e1006601 |doi=10.1371/journal.pgen.1006601 |issn=1553-7404 |pmc=5333797 |pmid=28253255 |doi-access=free}}</ref>

Over the course of their evolution, probobscideans grew in size. With that came longer limbs and wider feet with a more [[digitigrade]] stance, along with a larger head and shorter neck. The trunk evolved and grew longer to provide reach. The number of premolars, incisors, and canines decreased, and the cheek teeth (molars and premolars) became longer and more specialised. The incisors developed into tusks of different shapes and sizes.<ref name=evolution/> Several species of proboscideans became isolated on islands and experienced [[insular dwarfism]],<ref name=Sukumar31>Sukumar, pp. 31–33.</ref> some dramatically reducing in body size, such as the {{Cvt|1|m}} tall [[dwarf elephant]] species ''[[Palaeoloxodon falconeri]]''.<ref>{{Cite journal |last1=Romano |first1=M. |last2=Manucci |first2=F. |last3=Palombo |first3=M. R. |date=2021 |title=The smallest of the largest: new volumetric body mass estimate and in-vivo restoration of the dwarf elephant Palaeoloxodon ex gr. ''P. falconeri'' from Spinagallo Cave (Sicily) |journal=Historical Biology |volume=33 |issue=3 |pages=340–353 |doi=10.1080/08912963.2019.1617289 |bibcode=2021HBio...33..340R |s2cid=181855906}}</ref>

===Living species===
{| class="wikitable"
|+ style="text-align: centre;" |
! Name
! Size
! Appearance 
! Distribution
! Image
|-
| [[African bush elephant]] (''Loxodonta africana'')
| '''Male''': {{cvt|304|-|336|cm|ftin|0|abbr=on}} (shoulder height), {{cvt|5.2|-|6.9|MT|ST|1|abbr=on}} (weight); '''Female''': {{cvt|247|-|273|cm|ftin|0|abbr=on}} (shoulder height), {{cvt|2.6|-|3.5|MT|ST|1|abbr=on}} (weight).<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>
| Relatively large and triangular ears, concave back, diamond shaped molar ridges, wrinkled skin, sloping abdomen, and two finger-like extensions at the tip of the trunk.<ref name=Shoshani38>Shoshani, pp. 38–41.</ref>
| [[Sub-Saharan Africa]]; forests, savannahs, deserts, wetlands, and near lakes.<ref name=Shoshani42>Shoshani, pp. 42–51.</ref>
|[[File:African Elephant (Loxodonta africana) male (17289351322).jpg|200px]]
|-
| [[African forest elephant]] (''Loxodonta cyclotis'')
| {{cvt|209|-|231|cm|ftin|0|abbr=on}} (shoulder height), {{cvt|1.7|-|2.3|MT|ST|1|abbr=on}} (weight).<ref name="Larramendi, A. 2015"/> 
| Similar to the bush species, but with smaller and more rounded ears and thinner and straighter tusks.<ref name=Shoshani38/><ref name=Shoshani42 /> 
| [[West Africa|West]] and [[Central Africa]]; [[Atlantic Equatorial coastal forests|equatorial forests]], but occasionally [[gallery forest]]s and forest/grassland [[ecotone]]s.<ref name=Shoshani42 />
|[[File:Loxodontacyclotis.jpg|200px]]
|-
| [[Asian elephant]] (''Elephas maximus'')
| '''Male''': {{cvt|261|-|289|cm|ftin|0|abbr=on}} (shoulder height), {{cvt|3.5|-|4.6|MT|ST|1|abbr=on}} (weight); '''Female''': {{cvt|228|-|252|cm|ftin|0|abbr=on}} (shoulder height), {{cvt|2.3|-|3.1|MT|ST|1|abbr=on}} (weight).<ref name="Larramendi, A. 2015"/>
| Relatively small ears, convex or level back, dish-shaped forehead with two large bumps, narrow molar ridges, smooth skin with some blotches of [[depigmentation]], a straightened or saggy abdomen, and one extension at the tip of the trunk.<ref name=Shoshani38/>
|[[South Asia|South]] and [[Southeast Asia]]; habitats with a mix of grasses, low woody plants, and trees, including dry [[Deccan thorn scrub forests|thorn-scrub forests]] in southern India and Sri Lanka and [[evergreen forest]]s in [[Malay Peninsula|Malaya]].<ref name=Shoshani42/><ref name=Asian>{{cite journal|author1=Shoshani, J.|author2=Eisenberg, J. F.|year=1982|title=''Elephas maximus''|journal=Mammalian Species|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|access-date=27 October 2012|archive-date=24 September 2015|archive-url=https://web.archive.org/web/20150924121940/http://www.science.smith.edu/msi/pdf/i0076-3519-182-01-0001.pdf|url-status=dead}}</ref>
|[[File:Elephas maximus (Bandipur).jpg|200px]]
|}

==Anatomy==
[[File:Elephant skeleton.jpg|thumb|African bush elephant skeleton]]
Elephants are the largest living terrestrial animals. The skeleton 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> The skull contains air cavities ([[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. By contrast, the lower jaw is dense. The cranium is particularly large and provides enough room for the attachment of muscles to support the entire head.<ref name=Shoshani68>Shoshani, pp. 68–70.</ref> The skull is built to withstand great stress, particularly when fighting or using the tusks. The brain is surrounded by arches in the skull, which serve as protection.<ref>Kingdon, p. 11.</ref> Because of the size of the head, the neck is relatively short to provide better support.<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=480–487 |doi=10.1016/S0169-5347(98)01491-8 |pmid=21238404|bibcode=1998TEcoE..13..480S }}</ref>
Elephants are [[homeothermy|homeotherms]] and maintain their average body temperature at ~ 36&nbsp;°C (97&nbsp;°F), with a minimum of 35.2&nbsp;°C (95.4&nbsp;°F) during the cool season, and a maximum of 38.0&nbsp;°C (100.4&nbsp;°F) during the hot dry season.<ref>{{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|access-date=14 May 2021|archive-date=15 May 2021|archive-url=https://web.archive.org/web/20210515162738/https://link.springer.com/article/10.1007/s00360-018-1170-5|url-status=live|url-access=subscription}}</ref>

===Ears and eyes===
[[File:Angry elephant ears.jpg|thumb|African bush elephant with ears spread in a threat or attentive position and visible blood vessels]]
Elephant ear flaps, or [[pinna (anatomy)|pinnae]], are {{convert|1|–|2|mm|in|abbr=on}} thick in the middle with a thinner tip and supported by a thicker base. They contain numerous blood vessels called [[capillary|capillaries]]. Warm blood flows into the capillaries, releasing excess heat into the environment. This effect is increased by flapping the ears back and forth. 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 name=Shoshani68 /><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=638–647|doi=10.1007/s12045-008-0070-5}}</ref> The [[ossicles]] are adapted for hearing low frequencies, being most sensitive at 1 [[Hertz|kHz]].<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=1122–1123|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>

Lacking a [[lacrimal apparatus]] (tear duct), the eye relies on the [[harderian gland]] in the orbit to keep it moist. A durable [[nictitating membrane]] shields the globe. The animal's [[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 [[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=335–344|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 />

===Trunk===
{{Redirect|Elephant trunk}}
[[File:African elephant warning raised trunk.jpg|thumb|right|African bush elephant with its trunk raised, a behaviour often adopted when trumpeting]]
The elongated and [[Prehensility|prehensile]] trunk, or [[proboscis]], consists of both the nose and upper lip, which fuse in early [[Fetus|fetal]] development.<ref name=evolution /> This versatile appendage contains up to 150,000 separate [[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 [[Dorsoventral|dorsal, ventral]], and [[Lateral and medial|lateral]] muscles, while the latter are divided into [[Transverse plane|transverse]] and [[wikt:radiate|radiating]] muscles. The muscles of the trunk connect to a bony opening in the skull. The [[nasal septum]] consists of small elastic muscles between the nostrils, which are divided by [[cartilage]] at the base.<ref name=Shoshani74>Shoshani, pp. 74–77.</ref> A unique proboscis nerve – a combination of the [[maxillary nerve|maxillary]] and [[facial nerve]]s – lines each side of the appendage.<ref name=trunk />

As a [[muscular hydrostat]], the trunk moves through finely controlled muscle contractions, working both with and against each other.<ref name="trunk" /> Using three basic movements: bending, twisting, and longitudinal stretching or retracting, the trunk has near unlimited flexibility. Objects grasped by the end of the trunk can be moved to the mouth by curving the appendage inward. The trunk can also bend at different points by creating stiffened "pseudo-joints". The tip can be moved in a way similar to the human hand.<ref>{{cite journal|last1=Dagenais|first1=P|last2=Hensman|first2=S|last3=Haechler|first3=V|last4=Milinkovitch|first4=M. C.|year=2021|title=Elephants evolved strategies reducing the biomechanical complexity of their trunk|journal=Current Biology|volume=31|issue=21|pages=4727–4737|doi=10.1016/j.cub.2021.08.029|pmid=34428468|s2cid=237273086|doi-access=free|bibcode=2021CBio...31E4727D}}</ref> The skin is more elastic on the dorsal side of the elephant trunk than underneath; allowing the animal to stretch and coil while maintaining a strong grasp.<ref>{{cite journal|last1=Schulz|first1=A. K.|last2=Boyle|first2=M|last3=Boyle|first3=C|last4=Sordilla|first4=S|last5=Rincon|first5=C|last6=Hooper|first6=K|last7=Aubuchon|first7=C|last8=Reidenberg|first8=J. S.|last9=Higgins|first9=C|last10=Hu|first10=D. L.|year=2022|title=Skin wrinkles and folds enable asymmetric stretch in the elephant trunk|journal=Proceedings of the National Academy of Sciences|volume=119|issue=31|page=e2122563119|doi=10.1073/pnas.2122563119|doi-access=free |pmid=35858384 |pmc=9351381 |bibcode=2022PNAS..11922563S }}</ref> The flexibility of the trunk is aided by the numerous wrinkles in the skin.<ref>{{cite journal|last1=Schulz|first1=A. K.|last2=Kaufmann|first2=L. V.|last3=Reveyaz|first3=N|last4=Ritter|first4=C|last5=Hildebrandt|first5=T|last6=Brecht|first6=M|year=2024|title=Elephants develop wrinkles through both form and function|journal= Royal Society Open Science|volume=11|issue=10 |page=240851|doi=10.1098/rsos.240851|pmid=39386989 |pmc=11461087 |bibcode=2024RSOS...1140851S }}</ref> The African elephants have two finger-like extensions at the tip of the trunk that allow them to pluck small food. The Asian elephant has only one and relies more on wrapping around a food item.<ref name="Shoshani38" /> Asian elephant trunks have better [[motor coordination]].<ref name="Shoshani74" />
[[File:Asian Elephant, Royal Chitwan National Park.jpg|thumb|right|Asian elephant drinking water with trunk]]
The trunk's extreme flexibility allows it to forage and wrestle other elephants with it. It is powerful enough to lift up to {{convert|350|kg|lb|abbr=on}}, but it also has the precision to crack a peanut shell without breaking the seed. With its trunk, an elephant can reach items up to {{convert|7|m|ft|abbr=on}} high and dig for water in the mud or sand below. It also uses it to clean itself.<ref name=Kingdon9>Kingdon, p. 9.</ref> 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=371–379|doi=10.1037/0735-7036.117.4.371|pmid=14717638}}</ref> Elephant trunks are capable of powerful siphoning. They can expand their nostrils by 30%, leading to a 64% greater nasal volume, and can breathe in almost 30 times faster than a human sneeze, at over {{convert|150|m/s|abbr=on}}.<ref name="Schulz">{{cite journal|author=Schulz, A. K.|author2= Ning Wu, Jia|author3= Sara Ha, S. Y.|author4= 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}}</ref> They suck up water, which is squirted into the mouth or over the body.<ref name=evolution /><ref name="Schulz"/> The trunk of an adult Asian elephant is capable of retaining {{convert|8.5|L|gal|abbr=on}} of water.<ref name=Shoshani74 /> They will also sprinkle dust or grass on themselves.<ref name=evolution /> When underwater, the elephant uses its trunk as a [[Submarine snorkel|snorkel]].<ref name=snorkel />

The trunk also acts as a sense organ. Its sense of smell may be four times greater than a [[bloodhound]]'s nose.<ref name=Sukumar149 /> The [[infraorbital nerve]], which makes the trunk sensitive to touch, is thicker than both the [[Optic nerve|optic]] and [[Cochlear nerve|auditory]] nerves. [[Whiskers]] grow all along the trunk, and are particularly packed at the tip, where they contribute to its tactile sensitivity. Unlike those of many mammals, such as cats and rats, elephant whiskers do not move independently ("whisk") to sense the environment; the trunk itself must move to bring the whiskers into contact with nearby objects. Whiskers grow in rows along each side on the ventral surface of the trunk, which is thought to be essential in helping elephants balance objects there, whereas they are more evenly arranged on the dorsal surface. The number and patterns of whiskers are distinctly different between species.<ref>{{Cite journal |last1=Deiringer |first1=Nora |last2=Schneeweiß |first2=Undine |last3=Kaufmann |first3=Lena V. |last4=Eigen |first4=Lennart |last5=Speissegger |first5=Celina |last6=Gerhardt |first6=Ben |last7=Holtze |first7=Susanne |last8=Fritsch |first8=Guido |last9=Göritz |first9=Frank |last10=Becker |first10=Rolf |last11=Ochs |first11=Andreas |last12=Hildebrandt |first12=Thomas |last13=Brecht |first13=Michael |date=2023-06-08 |title=The functional anatomy of elephant trunk whiskers |journal=Communications Biology |language=en |volume=6 |issue=1 |page=591 |doi=10.1038/s42003-023-04945-5 |pmid=37291455 |pmc=10250425 |issn=2399-3642}}</ref>

Damaging the trunk would be detrimental to an elephant's survival,<ref name=evolution /> although in rare cases, individuals have survived with shortened ones. One trunkless elephant has been observed to graze using its lips with its hind legs in the air and balancing on its front knees.<ref name=Shoshani74 /> [[Floppy trunk syndrome]] is a condition of trunk [[paralysis]] recorded in African bush elephants and involves the degeneration of the [[Peripheral nervous system|peripheral nerves]] and muscles. The disorder has been linked to lead poisoning.<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|archive-date=17 May 2008|archive-url=https://web.archive.org/web/20080517112426/http://www.newscientist.com/article/mg13618470.700-lead-in-lake-blamed-for-floppy-trunks-.html|url-status=live}}</ref>

===Teeth===
{{multiple image
|align=right
|direction=vertical
|width=220
|image1=Loxodonta africana - Molar of an adult.JPG|caption1=Molar of an adult African 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 [[incisor]]s, known as the [[tusk]]s; 12 [[Deciduous teeth|deciduous]] [[premolar]]s; and 12 [[Molar (tooth)|molars]]. Unlike most mammals, teeth are not replaced by new ones emerging from the jaws vertically. Instead, new teeth start at the back of the mouth and 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. This is followed by four more tooth replacements at the ages of four to six, 9–15, 18–28, and finally in their early 40s. The final (usually sixth) set must last the elephant the rest of its life. Elephant teeth have loop-shaped dental ridges, which are 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 keep growing at about {{convert|17|cm|in|0|abbr=on}} a year. As the tusk develops, it is topped with smooth, cone-shaped [[tooth enamel|enamel]] that eventually wanes. The [[dentin]]e is known as [[ivory]] and has a [[Cross section (geometry)|cross-section]] of intersecting lines, known as "engine turning", which create diamond-shaped patterns. Being living tissue, tusks are fairly soft and about as dense as the mineral [[calcite]]. The tusk protrudes from a socket in the skull, and most of it is external. At least one-third of the tusk contains the [[Pulp (tooth)|pulp]], and some have nerves that stretch even further. Thus, it would be difficult to remove it without harming the animal. When removed, ivory will dry up and crack if not kept cool and wet. Tusks function in digging, debarking, marking, moving objects, and fighting.<ref name=Shoshani71 />

Elephants are usually right- or left-tusked, similar to humans, who are typically [[Handedness|right- or left-handed]]. The dominant, or "master" tusk, is typically more worn down, as it is shorter and blunter. For 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 more massive.<ref>Sukumar, p. 120</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 elephant]]s.<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|archive-date=21 November 2015|archive-url=https://web.archive.org/web/20151121185117/http://www.enn.com/wildlife/article/29620|url-status=live}}</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=dead|access-date=2021-10-28|website=[[National Geographic]]|language=en|archive-url=https://web.archive.org/web/20210303222242/https://www.nationalgeographic.com/animals/article/wildlife-watch-news-tuskless-elephants-behavior-change/ |archive-date=3 March 2021 }}</ref> has resulted in an effective [[selection pressure]] 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|bibcode=2015EcoEv...5.5216C }}</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|bibcode=1995AfJEc..33..230J }}</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:Elephant Skin (4526726581).jpg|thumb|Asian elephant skin]]
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, [[anus]], and inside of the ear is considerably thinner. Elephants are typically grey, but African elephants look brown or reddish after rolling in coloured mud. Asian elephants have some patches of depigmentation, particularly on the head. Calves have brownish or reddish hair, with the head and back being particularly hairy. As elephants mature, their hair darkens and becomes sparser, but dense concentrations of hair and bristles remain on the tip of the tail and parts of the head and genitals. 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 help 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 |page=e47018 |doi=10.1371/journal.pone.0047018|pmid=23071700 |pmc=3468452 |bibcode=2012PLoSO...747018M |doi-access=free }}</ref>

Although tough, an elephant's skin is very sensitive and requires [[Wallowing|mud baths]] to maintain moisture and protection from burning and insect bites. After bathing, the elephant will usually use its trunk to blow dust onto its body, which dries into a protective crust. Elephants have difficulty releasing heat through the skin because of their low [[surface-area-to-volume ratio]], which is many times smaller than that of a human. They have even been observed lifting up their legs to expose their soles to the air.<ref name=Shoshani66 /> Elephants only have [[sweat gland]]s between the toes,<ref>{{cite journal|last1=Lamps|first1=L. W.|last2=Smoller|first2=B. R.|last3=Rasmussen|first3=L. E. L.|last4=Slade|first4=B. E.|last5=Fritsch|first5=G|last6=Godwin|first6=T. E.|year=2001|title=Characterization of interdigital glands in the Asian elephant (Elephas maximus)|journal=Research in Veterinary Science|volume=71|issue=3|pages=197–200|doi=10.1053/rvsc.2001.0508|pmid=11798294 }}</ref> but the skin allows water to disperse and evaporate, cooling the animal.<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> In addition, cracks in the skin may reduce dehydration and allow for increased thermal regulation in the long term.<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>

=== Legs, locomotion, and posture ===
{{multiple image
|align=right
|direction=vertical
|width=220
|image1=Desert elephant (Loxodonta africana) feet.jpg|caption1=Front feet of bush elephant
|image2=Elephant Walking animated.gif|caption2=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]]s in place of [[medullary cavity|medullary cavities]]. This strengthens the bones while still allowing [[haematopoiesis]] (blood cell creation).<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=781–792|doi=10.1111/j.1469-7580.2006.00648.x|pmid=17118065|pmc=2048995}}</ref> The position of the limbs and leg bones allows an elephant to stand still for extended periods of time without tiring. Elephants are incapable of turning their [[Manus (anatomy)|manus]] as the [[ulna]] and [[Radius (bone)|radius]] of the front legs are secured in [[pronation]].<ref name=Shoshani69 /> Elephants may also lack the [[Pronator quadratus muscle|pronator quadratus]] and [[Pronator teres muscle|pronator teres]] muscles or have very small ones.<ref>Shoshani, p. 74.</ref> The circular feet of an elephant have soft tissues, or "cushion pads" beneath the manus or [[Pes (anatomy)|pes]], which allow them to bear the animal's great mass.<ref name=cushion /> They appear to have a [[Sesamoid bone|sesamoid]], an extra "toe" similar in placement to a [[giant panda]]'s extra "thumb", that also helps in weight distribution.<ref>{{cite journal|last1=Hutchinson|first1=J. R.|last2=Delmer|first2=C|last3=Miller|first3=C. E.|last4=Hildebrandt|first4=T|last5=Pitsillides|first5=A. A.|last6=Boyde|first6=A|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 |s2cid=206536505 |url=https://researchonline.rvc.ac.uk/id/eprint/5612/1/5612.pdf }}</ref> As many as five toenails can be found on both the front and hind feet.<ref name=Shoshani38 />

Elephants can move both forward and backward, but are incapable of [[trotting]], [[jumping]], or [[Horse gait|galloping]]. They can move on land only by walking or [[Ambling gait|ambling]]: a faster gait similar to running.<ref name=Shoshani69 /><ref name="kinetics"/> In walking, the legs act as pendulums, with the hips and shoulders moving up and down while the foot is planted on the ground. The fast gait does not meet all the criteria of running, since there is no point where all the feet are off the ground, although the elephant uses its legs much like other running animals, and can move faster by quickening its stride. 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}}. 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="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=3812–3827|pmid=16985198 |doi = 10.1242/jeb.02443|doi-access=free}}</ref><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=493–494 |doi=10.1038/422493a |pmid=12673241 |issue=6931 |bibcode=2003Natur.422..493H |url=http://researchonline.rvc.ac.uk/id/eprint/124/ |access-date=3 January 2023 |archive-date=9 August 2022 |archive-url=https://web.archive.org/web/20220809231054/https://researchonline.rvc.ac.uk/id/eprint/124/ |url-status=live }}</ref> 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 can swim for up to six hours while completely waterborne, moving at {{convert|2.1|km/h|0|abbr=on}} and traversing up to {{convert|48|km|mi|abbr=on}} continuously.<ref>Shoshani, p. 60.</ref>

=== Internal systems ===
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.<ref name=Shoshani78 /> It is the largest of all terrestrial mammals.<ref name="posse">{{cite book|author=O'Connell, Caitlin|title="Elephant Don: The Politics of a Pachyderm Posse"|date=20 July 2016|publisher=University of Chicago Press |isbn=9780226380056|url=https://books.google.com/books?id=2ZElEAAAQBAJ&q=Elephants+have+the+largest+temporal+lobe+relative+to+the+size+of+its+body+among+any+land+mammals&pg=PA102|access-date=16 September 2022|archive-date=21 March 2023|archive-url=https://web.archive.org/web/20230321084756/https://books.google.com/books?id=2ZElEAAAQBAJ&q=Elephants+have+the+largest+temporal+lobe+relative+to+the+size+of+its+body+among+any+land+mammals&pg=PA102|url-status=live}}</ref> While the elephant brain is larger overall, it is [[Encephalization quotient|proportionally smaller than the human brain]]. At birth, an elephant's brain already weighs 30–40% of its adult weight. The [[cerebrum]] and [[cerebellum]] are well developed, and the [[temporal lobe]]s are so large that they bulge out laterally.<ref name=Shoshani78 /> Their temporal lobes are proportionally larger than those of other animals, including humans.<ref name=posse /> The throat of an elephant appears to contain a pouch where it can store water for later use.<ref name=evolution /> The [[larynx]] of the elephant is the largest known among mammals. The [[vocal fold]]s are anchored close to the [[epiglottis]] base. When comparing an elephant's vocal folds to those of a human, an elephant's are proportionally longer, thicker, with a greater cross-sectional area. In addition, they are located further up the vocal tract with an acute slope.<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>
[[File:Em - Loxodonta africana heart - GMZ 2.jpg|thumb|right|African elephant heart in a jar]]
The heart of an elephant weighs {{convert|12|–|21|kg|lb|abbr=on}}. Its [[Heart#Location and shape|apex]] has two pointed ends, an unusual trait among mammals.<ref name=Shoshani78 /> In addition, the [[Ventricle (heart)|ventricles]] of the heart split towards the top, a trait also found in 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 |page=59}}</ref> When upright, the elephant's heart beats around 28 beats per minute and actually speeds up to 35 beats when it lies down.<ref name=Shoshani78/> The blood vessels are thick and wide and can hold up under high blood pressure.<ref name="Marshall Cavendish"/> The lungs are attached to the [[Thoracic diaphragm|diaphragm]], and breathing relies less on the expanding 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|doi-access=free}}</ref> Elephants breathe mostly with the trunk but also with the mouth. They have a [[hindgut fermentation]] system, and their large and small intestines together reach {{convert|35|m|ft|0|abbr=on}} in length. Less than half of an elephant's food intake gets digested, despite the process lasting a day.<ref name=Shoshani78 /> An elephant's bladder can store up to 18 litres of urine<ref>{{Cite journal |last1=Yang |first1=Patricia J. |last2=Pham |first2=Jonathan |last3=Choo |first3=Jerome |last4=Hu |first4=David L. |date=2014-08-19 |title=Duration of urination does not change with body size |journal=Proceedings of the National Academy of Sciences |language=en |volume=111 |issue=33 |pages=11932–11937 |doi=10.1073/pnas.1402289111 |doi-access=free |issn=0027-8424 |pmc=4143032 |pmid=24969420|bibcode=2014PNAS..11111932Y }}</ref> and its [[kidneys]] can produce more than 50 [[litre]]s of urine per day.<ref name="FowlerMikota2006" />

===Sex characteristics===
{{anchor|Sex organs}}{{multiple image|image1=Penis asiatischer Elefant.JPG|image2=Vulve d'éléphante.jpg|perrow=2|total_width=220px|footer=Asian elephant penis and vulva}}A male elephant's testes, like other [[Afrotheria]],<ref name="Sharma Lehmann Stuckas Funke ">{{cite journal | last1=Sharma | first1=Virag | last2=Lehmann | first2=Thomas | last3=Stuckas | first3=Heiko | last4=Funke | first4=Liane | last5=Hiller | first5=Michael | title=Loss of RXFP2 and INSL3 genes in Afrotheria shows that testicular descent is the ancestral condition in placental mammals | journal=PLOS Biology |  volume=16 | issue=6 | date=2018 | issn=1545-7885 | doi=10.1371/journal.pbio.2005293 | page=e2005293 | pmid=29953435 | pmc=6023123 | doi-access=free }}</ref> are internally located 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 = 517–536 | doi = 10.1530/jrf.0.0130517 | pmid = 6029179 | doi-access = free | access-date = 10 July 2017 | archive-date = 20 July 2018 | archive-url = https://web.archive.org/web/20180720144623/http://www.reproduction-online.org/content/13/3/517.full.pdf | url-status = live }}</ref> The [[penis]] can be as long as {{convert|100|cm|in|0|abbr=on}} with a {{convert|16|cm|in|0|abbr=on}} wide base. It curves to an 'S' when fully erect and has an [[External urethral orifice (male)|orifice]] shaped like a Y. The female's [[clitoris]] may be {{convert|40|cm|in|0|abbr=on}}. The [[vulva]] is found lower than in other herbivores, between the hind legs instead of under the tail. Determining pregnancy status can be difficult due to the animal's large belly. The female's [[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 [[Temporin|temporal gland]], located on 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 these secretions.<ref name=Sukumar149>Sukumar, p. 149.</ref>

==Behaviour and ecology==
{{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
}}
Elephants are [[Herbivore|herbivorous]] and will eat leaves, twigs, fruit, bark, grass, and roots. African elephants mostly [[Browsing (herbivory)|browse]], while Asian elephants mainly [[Grazing (behaviour)|graze]].<ref name=Shoshani42 /> They can eat as much as {{convert|300|kg|lb|abbr=on}} of food and drink {{convert|40|L|gal|abbr=on}} of water in a day. Elephants tend to stay near water sources.<ref name=Shoshani42 /><ref name=Shoshani124 /> They have morning, afternoon, and nighttime feeding sessions. At midday, elephants rest under trees and may doze off while standing. Sleeping occurs at night while the animal is lying down.<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=1264–1271|doi=10.1038/nature04285|pmid=16251951|pmc=8760626|bibcode=2005Natur.437.1264S}}</ref> Both males and family groups typically move no more than {{convert|20|km|mi|0|abbr=on}} a day, but distances as far as {{convert|180|km|mi|0|abbr=on}} have been recorded in the [[Etosha National Park|Etosha]] region of Namibia.<ref>Sukumar, p. 159.</ref> Elephants go on seasonal migrations in response to changes in environmental conditions.<ref>Sukumar, p. 174.</ref> In northern Botswana, they travel {{convert|325|km|mi|0|abbr=on}} to the [[Chobe River]] after the local waterholes dry up in late August.<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 [[keystone species]]. Their habit of uprooting trees and undergrowth can transform savannah into grasslands;<ref name=Shoshani226 /> smaller herbivores can access trees mowed down by elephants.<ref name=Shoshani124 /> When they dig for water during droughts, they create waterholes that can be used by other animals. When they use waterholes, they end up making them bigger.<ref name=Shoshani226 /> At [[Mount Elgon]], elephants dig through caves and pave the way for [[ungulate]]s, hyraxes, bats, birds, and insects.<ref name=Shoshani226 /> Elephants are important [[Seed dispersal|seed dispersers]]; African forest elephants consume and deposit many seeds over great distances, with either no effect or a positive effect on [[germination]].<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=542–553|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|access-date=22 November 2012|archive-date=18 June 2013|archive-url=https://web.archive.org/web/20130618142027/http://faculty.washington.edu/timbillo/Readings%20and%20documents/CO2%20and%20Forests%20readings/Campos%20Arceiz%202011%20Elephant%20seed%20dispersal.pdf|url-status=live}}</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 smaller [[Malayan 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=220–227|doi=10.1111/j.1744-7429.2011.00784.x|bibcode=2012Biotr..44..220C |hdl=10261/56573 |s2cid=83604698 }}</ref> Because most of the food elephants eat goes undigested, their dung can provide food for other animals, such as [[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, elephant numbers have threatened several species of small birds that depend on woodlands. Their weight causes the soil to compress, leading to [[surface runoff|runoff]] and [[erosion]].<ref name=Shoshani124>Eltringham, pp. 124–27.</ref>

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.<ref name=Shoshani124 /> The size of adult elephants makes them nearly invulnerable to [[predator]]s.<ref name=Asian /> Calves may be preyed on by [[lion]]s, [[spotted hyena]]s, and [[Lycaon pictus|wild dogs]] in Africa<ref name=African>{{cite journal|author1=Laursen, L.|author2=Bekoff, M.|year=1978|title=''Loxodonta africana''|journal=Mammalian Species|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|s2cid=253949585 |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 [[tiger]]s in Asia.<ref name=Asian /> The lions of [[Chobe National Park#Geography and ecosystems|Savuti]], Botswana, have adapted to hunting elephants, targeting calves, juveniles or even sub-adults.<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|bibcode=2006AfJEc..44..279J }}</ref> There are rare reports of adult Asian elephants falling prey to tigers.<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]] |access-date=21 June 2014 |archive-date=11 May 2016 |archive-url=https://web.archive.org/web/20160511041022/http://www.newindianexpress.com/cities/kochi/article103095.ece |url-status=dead }}</ref> Elephants tend to have high numbers of parasites, particularly [[nematode]]s, compared to many other mammals. This may be due to elephants being less vulnerable to predation; in other mammal species, individuals weakened by significant [[parasite load]]s are easily killed off by predators, removing them from the population.<ref>Sukumar, p. 121.</ref>

===Social organisation===
[[File:Serengeti Elefantenherde1.jpg|thumb|A family of African bush elephants]]
Elephants are generally [[gregarious]] animals. African bush elephants in particular have a complex, stratified social structure.<ref name="deSilva1"/> Female elephants spend their entire lives in tight-knit [[Matrilineality|matrilineal]] family groups.<ref name=Sukumar175/> {{Anchor|Matriarch}} They are led by the [[Matriarchy#Other_animals|matriarch]], who is often the eldest female.<ref>Sukumar, p. 186.</ref> 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 found that the death of the matriarch led to greater stress 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|access-date=30 October 2012|archive-date=24 November 2014|archive-url=https://web.archive.org/web/20141124052942/http://randd.defra.gov.uk/Document.aspx?Document=WC05007_7719_FRP.pdf|url-status=live}}</ref> When her tenure is over, the matriarch's eldest daughter takes her place instead of her sister (if present).<ref name=African /> One study found that younger matriarchs take potential threats less seriously.<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=3270–3276 |doi=10.1098/rspb.2011.0168 |url=http://www.lifesci.sussex.ac.uk/cmvcr/Publications_files/ElephantLeadersProofs.pdf |pmid=21411454 |pmc=3169024 |access-date=26 December 2012 |archive-date=29 April 2013 |archive-url=https://web.archive.org/web/20130429173959/http://www.lifesci.sussex.ac.uk/cmvcr/Publications_files/ElephantLeadersProofs.pdf |url-status=live }}</ref> Large family groups may split if they cannot be supported by local 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>

At [[Amboseli National Park]], Kenya, female groups may consist of around ten members, including four adults and their dependent offspring. Here, a cow's life involves interaction with those outside her group. Two separate families may associate and bond with each other, forming what are known as bond groups. During the dry season, elephant families may aggregate into clans. These may number around nine groups, in which clans do not form strong bonds but defend their dry-season ranges against other clans. The Amboseli elephant population is further divided into the "central" and "peripheral" subpopulations.<ref name=Sukumar175>Sukumar, pp. 172, 175–79.</ref>

Female Asian elephants tend to have more fluid social associations.<ref name="deSilva1">{{cite journal |last1=de Silva |first1=S. |last2=Wittemyer |first2=G. |year=2012 |title=A comparison of social organization in Asian elephants and African savannah elephants |journal=International Journal of Primatology |volume=  33|issue= 5|pages= 1125–1141|doi=10.1007/s10764-011-9564-1 |s2cid=17209753 }}</ref> In Sri Lanka, there appear to be stable family units or "herds" and larger, looser "groups". 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, with only one or two adult females and their offspring. A group containing more than two cows and their offspring is known as a "joint family". Malay elephant populations have even smaller family units and do not reach levels above a bond group. Groups of African forest elephants typically consist of one cow 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|Lone bull: Adult male elephants spend much of their time alone or in single-sex groups]]
Adult males live separate lives. As he matures, a bull associates more with outside males or even other families. At Amboseli, young males may be away from their families 80% of the time by 14–15 years of age. When males permanently leave, they either live alone or with other males. The former is typical of bulls in dense forests. A [[dominance hierarchy]] exists among males, whether they are social or solitary. Dominance depends on age, size, and sexual condition.<ref name=Sukumar179 /> Male elephants can be quite sociable when not competing for mates and form vast and fluid social networks.<ref>{{cite journal|last1=Goldenberg|first1=S. Z.|last2=de Silva|first2=S.|last3=Rasmussen|first3=H. B.|last4=Douglas-Hamilton|first4=I.|last5=Wittemyer|first5=G.|year=2014|title=Controlling for behavioural state reveals social dynamics among male African elephants, ''Loxodonta africana''|journal=Animal Behaviour|volume=95|pages=111–119|doi=10.1016/j.anbehav.2014.07.002|s2cid=53152412 }}</ref><ref>{{cite journal|last1=LaDue|first1=C. A.|last2=Vandercone|first2=R. P. G.|last3=Kiso|first3=W. K.|last4=Freeman|first4=E. W.|year=2022|title=Social behavior and group formation in male Asian elephants (Elephas maximus): the effects of age and musth in wild and zoo-housed animals|journal=Animals|volume=12|issue=9|page=1215|doi=10.3390/ani12091215 |pmid=35565641 |pmc=9100748 |doi-access=free }}</ref> Older bulls act as the leaders of these groups.<ref>{{cite journal|last1=Allen|first1=C. R. B.|last2=Brent|first2=L. J. N.|last3=Motsentwa|first3=T|last4=Weiss|first4=M. N.|last5=Croft|first5=D. P.|year=2020|title=Importance of old bulls: leaders and followers in collective movements of all-male groups in African savannah elephants (''Loxodonta africana'')|journal=Scientific Reports|volume=10|issue=1|page=13996|doi=10.1038/s41598-020-70682-y|pmid=32883968 |pmc=7471917 |bibcode=2020NatSR..1013996A }}</ref> The presence of older males appears to subdue the aggression and "deviant" behaviour of younger ones.<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=425–426|doi=10.1038/35044191|pmid=11100713|bibcode=2000Natur.408..425S}}</ref> The largest all-male groups can reach close to 150 individuals. Adult males and females come together to breed. Bulls will accompany family groups if a cow is in [[Estrous cycle|oestrous]].<ref name=Sukumar179>Sukumar, pp. 179–83.</ref>

===Sexual behaviour===
====Musth====
{{Main|Musth}}
[[File:2005-tusker-musth-crop.jpg|thumb|right|upright|Indian elephant bull in musth]]
Adult males enter a state of increased [[testosterone]] known as [[musth]]. In a population in southern India, males first enter musth at 15 years old, but it is not very intense until they are older than 25. At Amboseli, no bulls under 24 were found to be in musth, while half of those aged 25–35 and all those over 35 were. In some areas, there may be seasonal influences on the timing of musths. The main characteristic of a bull's musth is a fluid discharged from the [[Temporin|temporal gland]] that runs down the side of his face. Behaviours associated with musth include walking with a high and swinging head, nonsynchronous ear flapping, picking at the ground with the tusks, marking, rumbling, and urinating in the [[penile sheath|sheath]]. The length of this varies between males of different ages and conditions, lasting from days to months.<ref name=Sukumar100 />

Males become extremely aggressive during musth. Size is the determining factor in [[Agonistic behaviour|agonistic]] encounters when the individuals have the same condition. In contests between musth and non-musth individuals, musth bulls win the majority of the time, even when the non-musth bull is larger. A male may stop showing signs of musth when he encounters a musth male of higher rank. Those of equal rank tend to avoid each other. Agonistic encounters typically consist of threat displays, chases, and minor sparring. Rarely do they full-on fight.<ref name=Sukumar100>Sukumar, pp. 100–08.</ref> 

There is at least one documented case of [[Infanticide (zoology)|infanticide]] among Asian elephants at Dong Yai Wildlife Sanctuary, with the researchers describing it as most likely normal behaviour among aggressive musth elephants.<ref>{{cite journal |last1=Wanghongsa |first1=Sawai |last2=Boonkird |first2=Kalyanee |title=On the incident of infanticide in wild elephants |journal=Wildlife Yearbook |date=2006 |volume=7 |pages=111–119}}</ref>

====Mating====
[[File:Elephantsmating.jpg|thumb|African elephant bull mating with a member of a female group]]
Elephants are [[Polygyny in animals|polygynous]] breeders,<ref>Sukumar, p. 89.</ref> and most [[Copulation (zoology)|copulations]] occur during rainfall.<ref name="Sukumar, p. 262">Sukumar, p. 262.</ref> An oestrous cow uses [[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 [[flehmen response]], which requires him to collect a chemical sample with his trunk and taste it with the [[vomeronasal organ]] at the roof of the mouth.<ref>Sukumar, pp. 98–99.</ref> The oestrous cycle of a cow lasts 14–16 weeks, with the [[follicular phase]] lasting 4–6 weeks and the [[luteal phase]] lasting 8–10 weeks. While most mammals have one surge of [[luteinizing hormone]] during the follicular phase, elephants have two. The first (or anovulatory) surge, appears to change the female's scent, signaling to males that she is in heat, 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> Cows over 45–50 years of age are less fertile.<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 | access-date = 24 October 2018 | archive-date = 24 October 2018 | archive-url = https://web.archive.org/web/20181024231940/https://www.researchgate.net/publication/222028670 | url-status = live }}</ref> Most mate-guarding is done by musth males, and females seek them out, particularly older ones.<ref>Sukumar, p. 113.</ref> 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 for comfort.<ref name=Shoshani106 /> During copulation, the male rests his trunk on the female.<ref>Kingdon, p. 69.</ref> The penis is mobile enough to move without 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|year=2006|publisher=John Wiley & Sons|isbn=978-0-8138-0676-1|page=353|access-date=17 October 2020|archive-date=21 March 2023|archive-url=https://web.archive.org/web/20230321084756/https://books.google.com/books?id=oCpiZA61tyQC&q=elephant+penis+mating&pg=PA353|url-status=live}}</ref> Before mounting, it curves forward and upward. Copulation lasts about 45 seconds and does not involve [[pelvic thrust]]ing or an 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/263 263–266]|year=1991|publisher=University of California Press|isbn=978-0-520-08085-0}}</ref>

[[Homosexual behavior in animals#Elephants|Homosexual behaviour]] has been observed 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 engage in [[Animal sexual behaviour#Autoeroticism or masturbation|mutual masturbation]] 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/427 427–430]|isbn=978-1-4668-0927-7}}</ref>

===Birth and development===
{{multiple images
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|image1=African elephant, Dzanga Sangha, Central African Republic (18322370163).jpg|caption1=An African forest elephant mother with her calf
|image2=L'éléphanteau têtant sa mère.JPG|caption2=Asian elephant calf suckling}}
[[Gestation]] in elephants typically lasts between one and a half and two years and the female will not give birth again for at least four years.<ref>Sukumar, pp. 259–60.</ref> The relatively long pregnancy is supported by several [[corpus luteum]]s 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=3687–3696|pmid=22719030|doi=10.1098/rspb.2012.1038|pmc= 3415912}}</ref> Births tend to take place during the wet season.<ref name="Sukumar, p. 262"/> Typically, only a single young is born, but twins sometimes occur.<ref name="pregnancy"/> Calves are born roughly {{convert|85|cm|in|abbr=on}} tall and with a weight of around {{convert|120|kg|lb|abbr=on}}.<ref name=Shoshani106>Moss, pp. 106–13.</ref> They are [[precocial]] and quickly stand and walk to follow their mother and family herd.<ref name=Sukumar126 /> A newborn calf will attract the attention of all the 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 limits access to her young. [[Alloparenting]] – where a calf is cared for by someone other than its mother – takes place in some family groups. Allomothers are typically aged two to twelve years.<ref name=Shoshani106 />

For the first few days, the newborn is unsteady on its feet and needs its mother's help. It relies on touch, smell, and hearing, as its eyesight is less developed. With little coordination in its trunk, it can only flop it around which may cause it to trip. When it reaches its second week, the calf can walk with more balance and has more control over its trunk. After its first month, the trunk can grab and hold objects but still lacks sucking abilities, and the calf must bend down to drink. It continues to stay near its mother as it is still reliant on her. For its first three months, a calf relies entirely on its mother's milk, after which it begins to forage for vegetation and can use its trunk to collect water. At the same time, there is progress in lip and leg movements. By nine months, mouth, trunk, and foot coordination are mastered. Suckling bouts tend to last 2–4 min/hr for a calf younger than a year. After a year, a calf is fully capable of grooming, drinking, and feeding itself. It still needs its mother's milk and protection until it is at least two years old. Suckling after two years may improve growth, health, and fertility.<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|archive-date=29 September 2014|archive-url=https://web.archive.org/web/20140929112616/http://www.elephantsforever.co.za/life-cycle.html|url-status=live}}</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|archive-date=30 May 2014|archive-url=https://web.archive.org/web/20140530064627/http://www.elephantsforever.co.za/life-cycle-adolescence.html|url-status=live}}</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|archive-date=12 November 2012|archive-url=https://web.archive.org/web/20121112161638/http://news.bbc.co.uk/2/hi/asia-pacific/2800737.stm|url-status=live}}</ref>

===Communication===
<!-- Please do not add any more examples to this section. This subject already has its own article -->
{{main article|Elephant communication}}
{{multiple images
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|image1=Three elephant's curly kisses.jpg|caption1=Asian elephants greeting each other by inter-twining their trunks
|image2=Loxodonta africana oral rumble visualized with acoustic camera (25fps) - pone.0048907.s003.ogv|caption2=Low frequency rumble visualised with acoustic camera}}
Elephants communicate in various ways. Individuals greet one another by touching each other on the mouth, temporal glands, and genitals. This allows them to pick up chemical cues. Older elephants use trunk-slaps, kicks, and shoves to control younger ones. 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. A calf will press against its mother's front legs to signal it wants to rest and will touch her breast or leg when it wants to suckle.<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 tossing around dust and vegetation. They are usually bluffing when performing these actions. Excited elephants also raise their heads and spread their ears but additionally may raise their trunks. Submissive elephants will lower their heads and trunks, as well as flatten their ears against their necks, while those that are ready to fight will bend their ears in a V shape.<ref>Payne and Langbauer, pp. 119–20.</ref>

Elephants produce several vocalisations—some of which pass through the trunk<ref name=Shoshani120>Payne and Langbauer, pp. 120–21.</ref>—for both short and long range communication. This includes trumpeting, [[Bellow (sound)|bellowing]], [[roaring]], [[growling]], [[Bark (sound)|barking]], snorting, and [[Rumble (noise)|rumbling]].<ref name=Shoshani120/><ref>Sukumar, p. 141.</ref> Elephants can produce [[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 [[sound pressure]] levels of 85–90&nbsp;[[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|bibcode=1986BEcoS..18..297P }}</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, possibly over {{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=421–431|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 [[seismic communication|communicate with seismics]], vibrations produced by impacts on the earth's surface or acoustical waves that travel through it. An individual foot stomping 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 by rumbles 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=842–850 |doi=10.1007/s00265-005-0136-2 |url=http://www.sscnet.ucla.edu/anthro/bec/papers/OConnellRodwell_4.16.07.pdf |issue=6 |bibcode=2006BEcoS..59..842O |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=3066–3072|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 -->
{{Main|Elephant cognition}}
[[File:Insightful-Problem-Solving-in-an-Asian-Elephant-pone.0023251.s005.ogv|thumb|Elephant rolling a block to allow it to reach food]]
Elephants are among the most intelligent animals. They exhibit [[Mirror test|mirror self-recognition]], an indication of [[self-awareness]] and [[cognition]] that has also been demonstrated in some [[ape]]s and [[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=17053–17057|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 [[Tool use by animals|species known to use tools]]. An Asian elephant has been observed fine-tuning branches for use as [[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=839–847|doi=10.1006/anbe.2001.1815}}</ref> Tool modification by these animals is not as advanced as that of [[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]]s which give them long lasting memories of their environment on a wide scale. Individuals may be able to remember where their family members are located.<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|hdl=10023/1612|hdl-access=free}}</ref>

Scientists debate the extent to which elephants feel [[emotion]]. They are attracted to 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 chimpanzees and dolphins, a dying or dead elephant may elicit attention and aid from others, including those from other groups. This has been interpreted as expressing "concern";<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, the ''Oxford Companion to Animal Behaviour'' (1987) said 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|author-link= Jeffrey Moussaieff Masson|author2=Susan McCarthy |year= 1996|publisher= Vintage|isbn= 978-0-09-947891-1 |page= 272}}</ref>

==Conservation==
===Status===
{{see also|List of elephant species by population}}
[[File:Elefanti.jpg|thumb|A family of African forest elephants in the [[Dzanga-Sangha Special Reserve]] wetlands. This species is considered to be critically endangered.]]
African bush elephants were listed as [[Endangered]] by the [[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 [[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 population of at least 1.3&nbsp;million elephants, possibly as high as 3.0&nbsp;million. A decade later, the population was estimated to be 609,000; with 277,000 in Central Africa, 110,000 in Eastern Africa, 204,000 in Southern Africa, and 19,000 in Western Africa. The population of rainforest elephants was lower than anticipated, at around 214,000 individuals. Between 1977 and 1989, elephant populations declined by 74% in East Africa. After 1987, losses in elephant numbers hastened, 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 various, with unconfirmed 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> The IUCN estimated that total population in Africa is estimated at to 415,000 individuals for both species combined as of 2016.<ref>{{cite web|author1=Thouless, C. R.|author2= Dublin, H. T.|author3= Blanc, J. J.|author4= Skinner, D. P.|author5= Daniel, T. E.|author6= Taylor, R. D.|author7= Maisels, F.|author8= Frederick, H. L.|author9= Bouché, P.|year=2016|title=African Elephant Status Report 2016|publisher=IUCN African Elephant Database|access-date=2021-06-05|url=https://portals.iucn.org/library/sites/library/files/documents/SSC-OP-060_A.pdf|archive-date=27 June 2021|archive-url=https://web.archive.org/web/20210627110047/https://portals.iucn.org/library/sites/library/files/documents/SSC-OP-060_A.pdf|url-status=live}}</ref>

African elephants receive at least some legal protection in every country where they are found. Successful conservation efforts in certain areas have led to high population densities while failures have led to declines as high as 70% or more of the course of ten years. As of 2008, local numbers were controlled by contraception or [[Species translocation|translocation]]. Large-scale [[culling]]s stopped in the late 1980s and early 1990s. In 1989, the African elephant was listed under Appendix I by the [[CITES|Convention on International Trade in Endangered Species of Wild Fauna and Flora]] (CITES), making trade illegal. Appendix II status (which allows restricted trade) was given to elephants in Botswana, Namibia, and Zimbabwe in 1997 and South Africa in 2000. In some countries, [[Trophy hunting|sport hunting]] of the animals is legal; Botswana, Cameroon, Gabon, Mozambique, Namibia, South Africa, Tanzania, Zambia, and Zimbabwe have CITES export quotas for elephant trophies.<ref name=IUCN />

In 2020, the IUCN listed the Asian elephant as [[endangered]] due to the population declining by half over "the last three generations".<ref name="IUCN2">{{cite iucn |author1=Williams, C. |display-authors=etal |year=2020 |url=https://www.iucnredlist.org/species/7140/45818198 |title=''Elephas maximus'' |access-date=2020-06-03}}</ref> Asian elephants once ranged from [[Western Asia|Western]] to [[East Asia]] and south to [[Sumatra]].<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]] may have stabilised.<ref name=IUCN2 />

===Threats===
{{See also|Elephant ivory|Elephant meat}}
[[File:Ivory trade.jpg|thumb|upright|Men with elephant tusks at [[Dar es Salaam]], Tanzania, {{Circa|1900}}]]
The [[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 was comparable to that of gold.<ref name=Shoshani202>Martin, pp. 202–07</ref> The ivory trade contributed to the fall of the African elephant population 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|archive-date=26 September 2015|archive-url=https://web.archive.org/web/20150926142015/http://ngm.nationalgeographic.com/2012/10/ivory/christy-text|url-status=dead}}</ref> Ivory was banned internationally by CITES in 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 as badly affected.<ref name=Shoshani202 /> Zimbabwe, Botswana, Namibia, Zambia, and Malawi wanted to continue the ivory trade and were allowed to, since their local populations were healthy, but only if their supplies were from culled individuals or those that died of natural causes.<ref name=ivory />

The ban allowed the elephant to recover in parts of Africa.<ref name=Shoshani202 /> In February 2012, 650 elephants in [[Bouba Njida National Park]], Cameroon, were slaughtered 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|archive-date=25 September 2015|archive-url=https://web.archive.org/web/20150925125357/http://www.theguardian.com/environment/2013/mar/19/86-elephants-chad-poaching-massacre|url-status=live}}</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|archive-date=21 December 2019|archive-url=https://web.archive.org/web/20191221145628/https://www.theguardian.com/environment/2015/sep/26/china-and-us-agree-on-ivory-ban-in-bid-to-end-illegal-trade-globally|url-status=live}}</ref>

Other threats to elephants include [[habitat destruction]] and [[Habitat fragmentation|fragmentation]]. The Asian elephant lives in areas with some of the highest human populations and may be confined to small islands of forest among human-dominated landscapes. Elephants commonly trample and consume crops, which contributes to conflicts with humans, and both elephants and humans have died by the hundreds as a result. Mitigating these conflicts is important for conservation. One proposed solution is the protection of [[wildlife corridor]]s which give populations greater interconnectivity and space.<ref name=IUCN2 /> Chili pepper products as well as guarding with defense tools have been found to be effective in preventing crop-raiding by elephants. Less effective tactics include [[beehive fence|beehive]] and [[electric fences]].<ref>{{cite journal|last1=Montgomery|first1=R. A.|last2=Raupp|first2=J|last3=Mukhwana|first3=M|last4=Greenleaf|first4=A|last5=Mudumba|first5=T|last6=Muruthi|first6=P|year=2022|title=The efficacy of interventions to protect crops from raiding elephants|journal=Ambio|volume=51|issue=3|pages=716–727|doi=10.1007/s13280-021-01587-x|pmid=34173175 |pmc=8800974 |bibcode=2022Ambio..51..716M }}</ref>

==Human relations==
===Working animal===
{{further|Captive elephants}}
[[File:Elefant pune.jpg|thumb|Working elephant as transport]]
Elephants have been [[working animal]]s since at least the [[Indus Valley civilization]] over 4,000 years ago<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, the age range when they are both more trainable and can work for more years.<ref name=Shoshani149 /> They were [[Mela shikar|traditionally captured with traps and lassos]], but since 1950, [[sedative|tranquillisers]] have been used.<ref>Wylie, pp. 120–23.</ref> Individuals of the Asian species have often been trained as working animals. Asian elephants are used to carry and pull both objects and people in and out of areas as well as lead people in religious celebrations. They are valued over mechanised tools as they can perform the same tasks but in more difficult terrain, with strength, memory, and delicacy. Elephants can learn over 30 commands.<ref name="Shoshani149">McNeely, pp. 149–50.</ref> Musth bulls are difficult and dangerous to work with and so are chained up until their 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,1960|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 |archive-date=1 June 2015 |archive-url=https://web.archive.org/web/20150601060613/http://www.fao.org/docrep/005/ad031e/ad031e0g.htm |url-status=live }}</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=Fuller, Thomas |title=Unemployed, Myanmar's Elephants Grow Antsy, and Heavier |url=https://www.nytimes.com/2016/01/31/world/asia/myanmar-logging-elephants-unemployment.html |archive-url=https://ghostarchive.org/archive/20220101/https://www.nytimes.com/2016/01/31/world/asia/myanmar-logging-elephants-unemployment.html |archive-date=2022-01-01 |url-access=limited |work=[[The New York Times]] |date=30 January 2016 |access-date=31 January 2016 }}{{cbignore}}</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=Lohanan, Roger  |date=February 2001 |access-date=31 January 2016 |publisher=[[FAO]] |archive-date=4 April 2016 |archive-url=https://web.archive.org/web/20160404162525/http://www.fao.org/docrep/005/ad031e/ad031e0r.htm |url-status=live }}</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 Center|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 -->
{{Main|War elephant}}
{{See also|Execution by elephant}}
[[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 described in [[Sanskrit]] texts as far back as 1500 BC. From South Asia, the use of elephants in warfare spread west to Persia<ref name=Shoshani146 /> and east to Southeast Asia.<ref name=SEA /> The Persians used them during the [[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=487–489|publisher=Bloomsbury Academic |isbn=978-1-57607-770-2}}</ref> War elephants were also employed in the Mediterranean and North Africa throughout the [[classical antiquity|classical period]] since the reign of [[Ptolemy II Philadelphus|Ptolemy II]] in Egypt. The [[Carthage|Carthaginian]] general [[Hannibal]] famously took African elephants across the [[Alps]] during his war with the Romans and reached the [[Po Valley]] in 218 BC with all of them alive, but died of disease and combat a year later.<ref name="Shoshani146">Wylie (2000), pp. 146–48.</ref>

An elephant's head and sides were equipped with armour, the trunk may have had a sword tied to it and tusks were sometimes covered with sharpened iron or brass. Trained elephants would attack both humans and horses with their tusks. They might have grasped an enemy soldier with the trunk and tossed him to their [[mahout]], or pinned the soldier to the ground and speared him. Some shortcomings of war elephants included their great visibility, which made them easy to target, and limited maneuverability compared to horses. [[Alexander the Great]] achieved victory over armies with war elephants by having his soldiers injure the trunks and legs of the animals which caused them to panic and become uncontrollable.<ref name=Shoshani146 />

===Zoos and circuses===
{{further|Captive elephants}}
[[File:Flickr - …trialsanderrors - Terrific flights over ponderous elephants, poster for Forepaugh ^ Sells Brothers, ca. 1899.jpg|thumb|Circus poster, {{circa|1900}}]]
Elephants have traditionally been a major part of [[zoo]]s and [[circus]]es around the world. In circuses, they are trained to perform tricks. The most famous circus elephant was probably [[Jumbo]] (1861 – 15 September 1885), who was a major attraction in the [[Ringling Bros. and Barnum & Bailey Circus|Barnum & Bailey Circus]].<ref>Shoshani, pp. 168–69.</ref><ref name=Shoshani184/> 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, imports of the species almost stopped by the end of the 1980s. Subsequently, the US received many 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 allow easy access to the animals and provide fund and knowledge for preserving their natural habitats, as well as safekeeping for the species. Opponents claim that 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|archive-date=23 September 2015|archive-url=https://web.archive.org/web/20150923235332/http://www.elephants.com/media/MSNBC_2_28_05.htm|url-status=live}}</ref> Elephants have been recorded displaying [[Stereotypy (non-human)|stereotypical behaviours]] in the form of wobbling the body or head and pacing the same route both forwards and backwards. 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|archive-date=24 November 2014|archive-url=https://web.archive.org/web/20141124052942/http://randd.defra.gov.uk/Document.aspx?Document=WC05007_7719_FRP.pdf|url-status=live}}</ref> One study claims wild elephants in protected areas of Africa and Asia live more than twice as long as those in European zoos; the median lifespan of elephants in European zoos being 17 years. Other studies suggest that elephants in zoos live a similar lifespan as 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|archive-date=4 May 2015|archive-url=https://web.archive.org/web/20150504050854/http://news.nationalgeographic.com/news/2008/12/081211-zoo-elephants.html|url-status=dead}}</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|archive-date=5 September 2015|archive-url=https://web.archive.org/web/20150905065353/http://www.humanesociety.org/issues/circuses_entertainment/facts/circus_myths.html|url-status=dead}}</ref> In testimony to a US federal court in 2009, Barnum & Bailey Circus CEO [[Kenneth Jeffrey Feld|Kenneth Feld]] acknowledged that circus elephants are struck behind their ears, under their chins, and on their legs with metal-tipped prods, called [[Elephant goad|bull hooks]] or ankus. Feld stated that these practices are necessary to protect circus workers and acknowledged that an elephant trainer was rebuked for using an electric prod on an elephant. Despite this, he denied that any of these practices hurt the animals.<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 positive reinforcement when training his animals.<ref>Wylie, p. 142.</ref> 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|archive-date=22 September 2017|archive-url=https://web.archive.org/web/20170922050831/http://www.cnn.com/2016/05/01/us/ringling-bros-elephants-last-show/index.html|url-status=live}}</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 | archive-date = 20 September 2013 | archive-url = https://web.archive.org/web/20130920190617/http://www.smh.com.au/articles/2006/02/16/1140064206413.html | url-status = live }}</ref><ref>{{cite news|author=Siebert, C.|date=8 October 2006|title=An Elephant Crackup?|work=[[The New York Times]]|url=https://www.nytimes.com/2006/10/08/magazine/08elephant.html|access-date=25 October 2012|archive-date=14 November 2012|archive-url=https://web.archive.org/web/20121114151839/http://www.nytimes.com/2006/10/08/magazine/08elephant.html|url-status=live}}</ref> In parts of India, male elephants have entered villages at night, destroying homes and killing people. From 2000 to 2004, 300 people died in [[Jharkhand]], and in [[Assam]], 239 people were reportedly killed between 2001 and 2006.<ref name="huggler" />
Throughout the country, 1,500 people were killed by elephants between 2019 and 2022, which led to 300 elephants being killed in kind.<ref>{{Cite news |last=Krishnan |first=Murali |date=19 August 2022 |title=India sees uptick in deadly elephant attacks |work=[[Deutsche Welle]] |url=https://www.dw.com/en/india-sees-more-deadly-elephant-attacks-as-habitats-shrink/a-62863404 |access-date=11 June 2023}}</ref> Local people have reported that some elephants were drunk during the attacks, though officials have disputed this.<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 | archive-date = 26 May 2007 | archive-url = https://web.archive.org/web/20070526151446/http://news.bbc.co.uk/1/hi/world/south_asia/241781.stm | url-status = live }}</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 | archive-date = 26 December 2007 | archive-url = https://web.archive.org/web/20071226035744/http://news.bbc.co.uk/1/hi/world/south_asia/482001.stm | url-status = live }}</ref> Purportedly drunk elephants attacked an Indian village in December 2002, killing six people, which led to the retaliatory slaughter 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 | archive-date = 8 April 2013 | archive-url = https://web.archive.org/web/20130408031502/http://news.bbc.co.uk/2/hi/south_asia/2583891.stm | url-status = live }}</ref>

===Cultural significance===
<!-- Please do not add any more examples to this section. This subject already has its own article -->
{{Main|Cultural depictions of elephants}}
{{See also|Elephants in Kerala culture|List of elephants in mythology and religion|List of fictional pachyderms}}
Elephants have a universal presence in global culture. They have been represented in art since [[Paleolithic]] times. Africa, in particular, contains many examples of elephant [[rock art]], especially in the [[Sahara]] and southern Africa.<ref>Wylie, pp. 62–65.</ref> In Asia, the animals are depicted as [[Motif (visual arts)|motifs]] in [[Hinduism|Hindu]] and [[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 [[Ancient Rome|ancient Romans]], who kept the animals in captivity, depicted elephants more accurately than [[medieval]] Europeans who portrayed them more like fantasy creatures, with horse, bovine, and boar-like traits, and trumpet-like trunks. As Europeans gained more access to captive elephants during the 15th century, depictions of them became more accurate, including one made by [[Leonardo da Vinci]].<ref>Wylie, pp. 83–84.</ref>
[[File:Ganesha_asianartmuseumsf.jpg|thumb|[[Hinduism|Hindu]] god [[Ganesha]]]]
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. 158–65.</ref> Similar ideas existed among other African societies, who believed that their chiefs would be [[reincarnation|reincarnated]] as elephants. During the 10th century AD, the people of [[Igbo-Ukwu]], in modern-day Nigeria, placed elephant tusks underneath their dead leader's feet in the grave.<ref>Wylie, p. 79.</ref> The animals' importance is only [[Totemism|totemic]] in Africa but is much more significant in Asia.<ref>Sukumar, p. 87.</ref> 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 [[Ganesha]], is ranked equal with the supreme gods [[Shiva]], [[Vishnu]], and [[Brahma]] in some traditions.<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, but could also take these things away.<ref name="Shoshani158" /> In Buddhism, [[Gautama Buddha|Buddha]] is said to have taken the form of a [[White elephant (animal)|white elephant]] when he entered his [[Maya (mother of Buddha)|mother's]] womb to be reincarnated as a human.<ref>Sukumar, p. 62.</ref> 

In Western popular culture, elephants symbolise the exotic, especially since – as with the [[giraffe]], [[hippopotamus]], and [[rhinoceros]] – there are no similar animals familiar to Western audiences. As characters, elephants are most common in [[Children's literature|children's stories]], where they are portrayed positively. They are typically surrogates for humans with ideal human values. Many stories tell of isolated young elephants returning to or finding a family, such as "The Elephant's Child" from [[Rudyard Kipling]]'s ''[[Just So Stories]]'', [[The Walt Disney Company|Disney]]'s ''[[Dumbo]],'' and Kathryn and Byron Jackson's ''The Saggy Baggy Elephant''. Other elephant heroes [[Anthropomorphism|given human qualities]] include [[Jean de Brunhoff]]'s [[Babar the Elephant|Babar]], [[David McKee]]'s [[Elmer the Patchwork Elephant|Elmer]], and [[Dr. Seuss]]'s [[Horton the Elephant|Horton]].<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>

Several cultural references emphasise the elephant's size and strangeness. For instance, a "[[white elephant]]" is a byword for something that is weird, unwanted, and has no value.<ref name="Van Riper 73" /> The expression "[[elephant in the room]]" refers to something that is being ignored but ultimately must be addressed.<ref>Wylie, p. 90.</ref> The story of the [[blind men and an elephant]] involves blind men touching different parts of an elephant and trying to figure out what it is.<ref>Wylie, pp. 27–28.</ref>

==See also==
{{Portal|Mammals}}
* [[Animal track]]
* [[Desert elephant]]
* [[Elephants' graveyard]]
* [[List of individual elephants]] 
* [[Motty]], captive hybrid of an Asian and African elephant
* [[National Elephant Day (Thailand)]]
* [[World Elephant Day]]

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

==Bibliography==
{{Refbegin|60em}}
* {{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|access-date= 10 March 2017|archive-date= 21 March 2023|archive-url= https://web.archive.org/web/20230321084757/https://books.google.com/books?id=xdH_Gi92-fAC|url-status= live}}
* {{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=106–113}}
** {{cite book |author1=Payne, K. B. |author2=Langauer, W. B |chapter=Elephant Communication |title=Elephants: Majestic Creatures of the Wild |pages=116–123}}
** {{cite book |author=Eltringham, S. K. |chapter=Ecology and Behavior |title=Elephants: Majestic Creatures of the Wild |pages=124–127}}
** {{cite book |author=Wylie, K. C. |chapter=Elephants as War Machines |title=Elephants: Majestic Creatures of the Wild |pages=146–148}}
** {{cite book |author=McNeely, J. A. |chapter=Elephants as Beasts of Burden |title=Elephants: Majestic Creatures of the Wild |pages=149–150}}
** {{cite book |author=Smith, K. H. |chapter=The Elephant Domestication Centre of Africa |title=Elephants: Majestic Creatures of the Wild |pages=152–154}}
** {{cite book |author=McNeely, J. A. |chapter=Elephants in Folklore, Religion and Art |title=Elephants: Majestic Creatures of the Wild |pages=158–165}}
** {{cite book |author=Shoshani, S. L. |chapter=Famous Elephants |title=Elephants: Majestic Creatures of the Wild |pages=168–171}}
** {{cite book |author=Daniel, J. C. |chapter=The Asian Elephant Population Today |title=Elephants: Majestic Creatures of the Wild |pages=174–177}}
** {{cite book |author=Douglas-Hamilton, I. |chapter=The African Elephant Population Today |title=Elephants: Majestic Creatures of the Wild |pages=178–183}}
** {{cite book |author=Tuttle, C. D. |chapter=Elephants in Captivity |title=Elephants: Majestic Creatures of the Wild |pages=184–193}}
** {{cite book |author=Martin, E. B. |chapter=The Rise and Fall of the Ivory Market |title=Elephants: Majestic Creatures of the Wild |pages=202–207}}
** {{cite book |author=Shoshani, J. |chapter=Why Save Elephants? |title=Elephants: Majestic Creatures of the Wild |pages=226–229}}
* {{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= 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|access-date= 10 March 2017|archive-date= 21 March 2023|archive-url= https://web.archive.org/web/20230321084840/https://books.google.com/books?id=Aj75snWdfv8C|url-status= live}}
{{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}}
* {{Cite book |last=Nance |first=Susan |year=2013 |title=Entertaining Elephants: Animal Agency and the Business of the American Circus |location=Baltimore, MD |publisher=Johns Hopkins University Press}}
* [[John Godfrey Saxe|Saxe, John Godfrey]] (1872). [[s:The Blindmen and the Elephant|"The Blindmen and the Elephant"]] at [[Wikisource]]. ''The Poems of John Godfrey Saxe''.
* {{cite book | last=Williams | first=Heathcote |author-link=Heathcote Williams | title=Sacred Elephant | location=New York | publisher=Harmony Books | year=1989 | isbn=978-0-517-57320-4}}

== External links ==
{{Sister project links |wikt=elephant |commons=Category:Elephants |n=Category:Elephants |q=Elephants |voy=Elephants }}
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* [https://elephantconservation.org International Elephant Foundation]

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