View source for African elephant

{{Short description|Genus comprising two living elephant species}}
{{Other uses}}
{{Use dmy dates|date=September 2020}}
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{{Automatic taxobox
| name = African elephants
| fossil_range = Late [[Miocene]] – [[Holocene|Present]], {{fossilrange|7|0}}
| image = African Elephant (Loxodonta africana) male (17289351322).jpg
| image_caption = [[African bush elephant]] bull in Kruger National Park
| image2 = Loxodontacyclotis.jpg
| image2_caption = [[African forest elephant]] in [[Nouabalé-Ndoki National Park]]
| taxon = Loxodonta
| authority = Anonymous, 1827<ref name=MSW3>{{MSW3 Proboscidea |id=11500008 |page=91 |heading=Genus ''Loxodonta''}}</ref>
| type_species = ''[[Loxodonta africana|Elephas africana]]''<ref name=MSW3/>
| type_species_authority = Blumenbach, 1797
| subdivision_ranks = Species and Subspecies
| subdivision = *''[[Loxodonta africana|L. africana]]''
*''[[Loxodonta cyclotis|L. cyclotis]]''
*[[Extinction|†]] ''[[Loxodonta adaurora|L. adaurora]]''
::† ''L. a. adaurora''
::† ''L. a. kararae''
*† ''[[Loxodonta atlantica|L. atlantica]]''
::† ''L. a. angammensis''
::† ''L. a. atlantica''
*† ''[[Loxodonta exoptata|L. exoptata]]''
*† ''[[Loxodonta cookei|L. cookei]]''
| range_map = African Elephant distribution map without borders.svg
| range_map_caption = {{legend|#008000|&nbsp;Range of living ''Loxodonta'' (2007)}}
}}

'''African elephants''' are members of the [[Genus (biology)|genus]] '''''Loxodonta''''' comprising two living [[elephant]] [[species]], the [[African bush elephant]] (''L. africana'') and the smaller [[African forest elephant]] (''L. cyclotis''). Both are social [[herbivore]]s with grey [[skin]]. However, they differ in the size and colour of their [[tusk]]s as well as the shape and size of their [[ear]]s and [[skull]]s.

Both species are at a pertinent risk of extinction according to the [[IUCN Red List]]; as of 2021, the bush elephant is considered [[Endangered species|endangered]] while the forest elephant is considered [[critically endangered]]. They are threatened by [[habitat loss]] and fragmentation, along with [[poaching]] for the illegal [[ivory trade]] in several range countries.

''Loxodonta'' is one of two extant genera in the family [[Elephantidae]]. The name refers to the lozenge-shaped enamel of their molar teeth. [[Fossil]] remains of ''Loxodonta'' species have been found in [[Africa]], spanning from the Late [[Miocene]] (from around 7–6 million years ago) onwards.

== Etymology ==
The name ''Loxodonta'' comes from the [[Ancient Greek]] words λοξός (''loxós, ''"slanting", "crosswise") and ὀδούς (''odoús'', "tooth"), referring to the [[Lozenge (shape)|lozenge]]-shaped [[Enamel of teeth|enamel]] of the [[molar teeth]], which differs significantly from the rounded shape of the [[Asian elephant]]'s molar enamel.<ref name="Cuvier">{{cite book |last1=Cuvier |first1=F. |title=Histoire Naturelle des Mammifères, avec des figures originales, coloriées, dessinées d'après des animaux vivans |publisher=A. Belain |year=1825 |editor1=Geoffroy-Saint-Hilaire, É. |volume=Tome 6 |location=Paris |pages=117–118 |chapter=Éléphants d'Afrique |editor2=Cuvier, F. |chapter-url=https://archive.org/details/histoirenaturel6geof/page/n115}}</ref>

== Taxonomy and evolution==
[[File:Loxodonta africana-cyclotis skulls PZSL.png|150px|thumb|Comparison of bush (left) and forest (right) elephant skulls in frontal view. Note the shorter and wider head of ''L. cyclotis'', with a concave instead of convex forehead.|left]]
[[File:African bush elephant skeleton.jpg|150px|thumb|A female African bush elephant skeleton on display at the Museum of Osteology, Oklahoma City|left]]
The first [[scientific description]] of the African elephant was written in 1797 by [[Johann Friedrich Blumenbach]], who proposed the [[scientific name]] ''Elephas africanus''.<ref>{{cite book |author=Blumenbach, J. F. |year=1797 |title=Handbuch der Naturgeschichte |trans-title=Handbook of Natural History |edition=Fifth |location=Göttingen |publisher=Johann Christian Dieterich |chapter=2. ''Africanus'' |page=125 |chapter-url=http://www.deutschestextarchiv.de/book/view/blumenbach_naturgeschichte_1797?p=147}}</ref>
''[[wikt:Loxodonta|Loxodonte]]'' was proposed as a [[Genus (biology)|generic]] name for the African elephant by [[Frédéric Cuvier]] in 1825. 
An anonymous author used the Latinized spelling ''Loxodonta'' in 1827.<ref>{{cite journal |author=Anonymous |year=1827 |title=Analytical Notices of Books. Histoire Naturelle des Mammifères, avec des Figures originale, dessinées d'après des Animaux vivans; &c. Par MM. Geoffroy-Saint-Hilaire, et F. Cuvier. Livraison 52 et 53 |journal=The Zoological Journal |volume=3 |issue=9 |pages=140–143 |url=https://archive.org/details/zoologicaljourna318271828sowe/page/140}}</ref> This author was recognized as authority by the [[International Code of Zoological Nomenclature]] in 1999.<ref name=MSW3 />

''Elephas'' (''Loxodonta'') ''cyclotis'' was proposed by [[Paul Matschie]] in 1900, who described three African elephant [[zoological specimen]]s from Cameroon whose [[skull]]s differed in shape from those of  elephant skulls collected elsewhere in Africa.<ref>{{cite journal |author=Matschie, P. |year=1900 |title=Geographische Abarten des Afrikanischen Elefanten |journal=Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin |volume=3 |pages=189–197 |url=https://archive.org/details/sitzungsberichte1900gese/page/194}}</ref> In 1936, [[Glover Morrill Allen]] considered this elephant to be a distinct species and called it the  'forest elephant';<ref>{{cite journal |last1=Allen |first1=G. M. |year=1936 |title=Zoological results of the George Vanderbilt African Expedition of 1934. Part II — The forest elephant of Africa |journal=Proceedings of the Academy of Natural Sciences of Philadelphia |volume=88 |pages=15–44 |jstor=4064188}}</ref> later authors considered it to be a [[subspecies]].<ref name=Laurson1978>{{cite journal |last1=Laurson |first1=B. |last2=Bekoff |first2=M. |title=''Loxodonta africana'' |journal=Mammalian Species |date=1978 |issue=92 |pages=1–8 |doi=10.2307/3503889 |jstor=3503889 |s2cid=253949585 |url=http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-092-01-0001.pdf |access-date=5 August 2010}}</ref><ref name=estes>{{cite book |last=Estes |first=R. D. |title=The Safari Companion: A Guide to Watching African Mammals Including Hoofed Mammals, Carnivores, and Primates |publisher=Chelsea Green Publishing Company |location=Vermont |year=1999 |edition=Revised and expanded |isbn=1-890132-44-6 |chapter=Elephant ''Loxodonta africana'' Family Elephantidae, Order Proboscidea |pages=223–233 |chapter-url=https://books.google.com/books?id=Xqp7poFviNcC&pg=PA223}}</ref> Morphological and genetic analyses have since provided evidence for species-level differences between the African bush elephant and the African forest elephant.<ref name=Grubb_al2000>{{Cite journal |author1=Grubb, P. |author2=Groves, C. P. |author3=Dudley, J. P. |author4=Shoshani, J. |name-list-style=amp |year=2000 |title=Living African elephants belong to two species: ''Loxodonta africana'' (Blumenbach, 1797) and ''Loxodonta cyclotis'' (Matschie, 1900) |journal=Elephant |volume=2 |issue=4 |pages=1–4 |doi=10.22237/elephant/1521732169|doi-access=free }}</ref><ref>{{cite journal |last1=Roca |first1=A. L. |last2=Georgiadis |first2=N. |name-list-style=amp |last3=Pecon-Slattery |first3=J. |last4=O'Brien |first4=S. J. |date=2001 |title=Genetic Evidence for Two Species of Elephant in Africa |journal=Science |volume=293 |issue=5534 |pages=1473–1477 |doi=10.1126/science.1059936 |pmid=11520983|bibcode=2001Sci...293.1473R |s2cid=38285623 |url=https://zenodo.org/record/1230804 }}</ref><ref name=RohlandMalaspinas2007>{{cite journal |last1=Rohland |first1=N. |last2=Malaspinas |first2=A.-S. |last3=Pollack |first3=J. L. |last4=Slatkin |first4=M. |last5=Matheus |first5=P. |last6=Hofreiter |first6=M. |name-list-style=amp |title=Proboscidean Mitogenomics: Chronology and Mode of Elephant Evolution Using Mastodon as Outgroup |journal=PLOS Biology |volume=5 |issue=8 |year=2007 |pages=e207 |doi=10.1371/journal.pbio.0050207 |pmid=17676977 |pmc=1925134 |doi-access=free }}</ref><ref name=Rohland2010>{{cite journal |last1=Rohland |first1=N. |last2=Reich |first2=D. |last3=Mallick |first3=S. |last4=Meyer |first4=M. |last5=Green |first5=R. E. |last6=Georgiadis |first6=N. J. |last7=Roca |first7=A. L. |last8=Hofreiter |first8=M. |name-list-style=amp |title=Genomic DNA sequences from mastodon and woolly mammoth reveal deep speciation of forest and savanna elephants |journal=[[PLOS Biology]] |volume=8 |issue=12 |pages=e1000564 |year=2010 |doi=10.1371/journal.pbio.1000564 |pmid=21203580 |pmc=3006346 |doi-access=free }}</ref><ref name=MurphyIshida2011>{{cite journal |last1=Murphy |first1=W. J. |last2=Ishida |first2=Y. |last3=Oleksyk |first3=T. K. |last4=Georgiadis |first4=N. J. |last5=David |first5=V. A. |last6=Zhao |first6=K. |last7=Stephens |first7=R. M. |last8=Kolokotronis |first8=S.-O. |last9=Roca |first9=A. L.|name-list-style=amp |title=Reconciling Apparent Conflicts between Mitochondrial and Nuclear Phylogenies in African Elephants |journal=PLOS ONE |volume=6 |issue=6 |year=2011 |pages=e20642 |doi=10.1371/journal.pone.0020642 |pmid=21701575 |pmc=3110795 |bibcode=2011PLoSO...620642I |doi-access=free}}</ref>

In 1907, [[Richard Lydekker]] proposed six African elephant subspecies based on the different sizes and shapes of their ears.<ref>{{cite journal |author=Lydekker, R. |year=1907 |title=The Ears as a Race-Character in the African Elephant |journal=Proceedings of the Zoological Society of London |issue=January to April |pages=380–403 |doi=10.1111/j.1096-3642.1907.tb01824.x |url=https://archive.org/details/proceedingsofzoo19071446zool/page/380}}</ref> They are all considered [[synonym (taxonomy)|synonym]]ous with the African bush elephant.<ref name=MSW3/>

A third species, the West African elephant, has also been proposed but  needs confirmation. It is thought that this lineage has been isolated from the others for 2.4 million years.<ref>{{cite journal |author1=Eggert, L.S. |name-list-style=amp |author2=Rasner, C.A. |author3=Woodruff, D.S. |year=2002 |journal=Proceedings of the Royal Society of London. Series B: Biological Sciences |volume=269 |issue=1504 |pages=1993–2006 |title=The evolution and phylogeography of the African elephant inferred from mitochondrial DNA sequence and nuclear microsatellite markers |doi=10.1098/rspb.2002.2070|pmid=12396498 |pmc=1691127 }}</ref>

=== Extinct African elephants ===
Between the late 18th and 21st centuries, the following extinct African elephants were described on the basis of [[fossil]] remains:
*[[North African elephant]] ({{extinct}} ''Loxodonta africana pharaohensis'') proposed by [[Paulus Edward Pieris Deraniyagala]] in 1948 was a specimen from [[Fayum]] in Egypt.<ref>{{cite book |author=Deraniyagala, P. E. P. |year=1955 |title=Some extinct elephants, their relatives, and the two living species |publisher=Ceylon National Museums Publication |location=Colombo}}</ref>
* {{extinct}} ''[[Loxodonta atlantica]]'' was proposed as ''Elephas atlanticus'' by [[Auguste Pomel]] in 1879 based on a [[skull]] and bones found in [[Tighennif|Ternifine]], Algeria.<ref>{{cite book |last1=Pomel |first1=A. |year=1897 |series=Paléontologie : monographies |title=Les éléphants quaternaires |location=Alger |publisher=P. Fontana & Co. |pages=42–59 |chapter=''Elephas atlanticus'' Pom. |chapter-url=https://gallica.bnf.fr/ark:/12148/bpt6k62754098/f48}}</ref>
* {{extinct}} ''[[Loxodonta exoptata]]'' proposed by [[:de:Wilhelm Otto Dietrich|Wilhelm Otto Dietrich]] in 1941 was based on teeth found in [[Laetoli]], [[Tanzania]].<ref>{{cite journal |last1=Dietrich |first1=W. O. |year=1941 |title=Die säugetierpaläontologischen Ergebnisse der Kohl-Larsen'schen Expedition 1937–1939 im nördlichen Deutsch-Ostafrika |journal=Zentralblatt für Mineralogie, Geologie und Paläontologie |volume=B |issue=8 |pages=217–223}}</ref>
* {{extinct}} ''[[Loxodonta adaurora]]'' proposed by [[Vincent Maglio]] in 1970 was a complete [[skeleton]] found in [[Kanapoi]], Kenya.<ref>{{cite journal |last1=Maglio |first1=V. J. |year=1970 |title=Four new species of Elephantidae from the Plio-Pleistocene of northwestern Kenya |journal=Breviora |issue=341 |pages=1–43 |url=https://archive.org/details/breviora326350harv/page/n291}}</ref>
* {{extinct}} ''[[Loxodonta cookei]]'' proposed by [[William J. Sanders]] in 2007 based on teeth found in the Varswater Formation at Langebaanweg, South Africa.<ref name = "Sanders, 2007">{{cite journal |last1=Sanders |first1=W. |date=2007 |title=Taxonomic review of fossil Proboscidea (Mammalia) from Langebaanweg, South Africa|journal=Transactions of the Royal Society of South Africa |volume=62 |issue=1 |edition=Online |doi=10.1080/00359190709519192| url = https://sites.lsa.umich.edu/billsanders/wp-content/uploads/sites/382/2016/04/AP-SANDERS-REVISED.pdf | pages=1–16|bibcode=2007TRSSA..62....1S |s2cid=27499106 }}</ref>

=== Phylogeny and evolution ===
Relationships of living and extinct elephants based on DNA, after Palkopoulou et al. 2018.<ref name=":0" />{{clade
|label1=[[Elephantidae]]
|1={{clade
|1={{clade
|1='''''Loxodonta''''' (African elephants) [[File:Elephas africanus - 1700-1880 - Print - Iconographia Zoologica - (white background).jpg|60 px]]
|2={{extinct}}''[[Palaeoloxodon]]'' (straight-tusked elephants)  [[File:Palaeoloxodon namadicus-bpk (cropped).jpg|60 px]]
}}
|2={{clade
                     |1=''[[Elephas]]'' (Asian elephants) [[File:Indian elephant white background.jpg|60 px]]
                     |2={{extinct}}''[[Mammuthus]]'' (mammoths) [[File:Mammuthus trogontherii122DB.jpg|70 px]]
                     }}}}
               }}The oldest species of ''Loxodonta'' known is ''[[Loxodonta cookei]],'' with remains of the species known from around 7–5 million years ago, from remains found in Chad, Kenya, Uganda, and South Africa.''<ref name=":2">{{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, 205–208 |language=en |doi=10.1201/b20016}}</ref>''

Analysis of nuclear [[DNA sequences]] indicates that the [[genetic divergence]] between African bush and forest elephants dates 2.6 – 5.6 million years ago. The African forest elephant was found to have a high degree of [[genetic diversity]], likely reflecting periodic fragmentation of their habitat during the  changes in the [[Pleistocene]].<ref name=Rohland2010/>

[[Gene flow]] between the two African elephant species was examined at 21 locations. The analysis revealed that several African bush elephants carried [[mitochondrial DNA]] of African forest elephants, indicating they [[Hybridisation (biology)|hybridised]] in the savanna-forest transition zone in ancient times.<ref name="Roca2004">{{cite journal |last1=Roca |first1=A. L. |last2=Georgiadis |first2=N. |last3=O'Brien |first3=S. J. |author3-link=Stephen J. O'Brien |name-list-style=amp |date=2004 |title=Cytonuclear genomic dissociation in African elephant species |journal=Nature Genetics |volume=37 |issue=1 |pages=96–100 |doi=10.1038/ng1485 |pmid=15592471 |s2cid=10029589 |url=http://planet.uwc.ac.za/nisl/Biodiversity/pdf/Nature%20Genet%202005%20Roca%20et%20al%20elephant%20phylgeogr.pdf |access-date=20 June 2019 |archive-date=27 November 2013 |archive-url=https://web.archive.org/web/20131127012455/http://planet.uwc.ac.za/nisl/Biodiversity/pdf/Nature%20Genet%202005%20Roca%20et%20al%20elephant%20phylgeogr.pdf |url-status=dead }}</ref> However, despite the hybridisation at the contact zone between the two species, there appears to have been little effective gene flow between the two species since their initial split.<ref name=":0" />
[[File:Palaeoloxodon_phylogeny.svg|thumb|243x243px|Phylogeny showing the placement of [[straight-tusked elephant]] (''[[Palaeoloxodon]] antiquus'') in relation to other elephantids based on nuclear genomes, after Palkopoulou et al. 2018, showing [[introgression]] from [[African forest elephant]]s]]
DNA from the European [[straight-tusked elephant]] (''Palaeoloxodon antiquus'') indicates that the extinct elephant genus ''[[Palaeoloxodon]]'' is more closely related to African elephants than to Asian elephants or mammoths. Analysis of the genome of ''P. antiquus'' also shows that ''Palaeloxodon'' extensively hybridised with African forest elephants, with the mitochondrial genome and over 30% of the nuclear genome  of ''P. antiquus'' deriving from ''L. cyclotis.'' This ancestry is closer to modern west African populations than to central African populations of forest elephants.<ref name=":0">{{Cite journal |author1=Eleftheria Palkopoulou |author2=Mark Lipson |author3=Swapan Mallick |author4=Svend Nielsen |author5=Nadin Rohland |author6=Sina Baleka |author7=Emil Karpinski |author8=Atma M. Ivancevic |author9=Thu-Hien To |author10=R. Daniel Kortschak |author11=Joy M. Raison |author12=Zhipeng Qu |author13=Tat-Jun Chin |author14=Kurt W. Alt |author15=Stefan Claesson |year=2018 |title=A comprehensive genomic history of extinct and living elephants |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=115 |issue=11 |pages=E2566–E2574 |bibcode=2018PNAS..115E2566P |doi=10.1073/pnas.1720554115 |pmc=5856550 |pmid=29483247 |doi-access=free |author16=Love Dalén |author17=Ross D. E. MacPhee |author18=Harald Meller |author19=Alfred L. Roca |author20=Oliver A. Ryder |author21=David Heiman |author22=Sarah Young |author23=Matthew Breen |author24=Christina Williams |author25=Bronwen L. Aken |author26=Magali Ruffier |author27=Elinor Karlsson |author28=Jeremy Johnson |author29=Federica Di Palma |author30=Jessica Alfoldi |author31=David L. Adelson |author32=Thomas Mailund |author33=Kasper Munch |author34=Kerstin Lindblad-Toh |author35=Michael Hofreiter |author36=Hendrik Poinar |author37=David Reich}}</ref> Analysis of Chinese ''Palaeoloxodon'' mitogenomes suggests that this forest elephant ancestry was widely shared among ''Palaeoloxodon'' species.<ref name=":4">{{Cite journal |last1=Lin |first1=Haifeng |last2=Hu |first2=Jiaming |last3=Baleka |first3=Sina |last4=Yuan |first4=Junxia |last5=Chen |first5=Xi |last6=Xiao |first6=Bo |last7=Song |first7=Shiwen |last8=Du |first8=Zhicheng |last9=Lai |first9=Xulong |last10=Hofreiter |first10=Michael |last11=Sheng |first11=Guilian |date=July 2023 |title=A genetic glimpse of the Chinese straight-tusked elephants |journal=Biology Letters |language=en |volume=19 |issue=7 |doi=10.1098/rsbl.2023.0078 |issn=1744-957X |pmc=10353889 |pmid=37463654 }}</ref>

== Description ==

=== Skin, ears, and trunk ===
[[File:African Bush Elephant Skull.jpg|thumb|A male African bush elephant skull on display at the [[Museum of Osteology]]]]
African elephants have grey folded skin up to {{convert|30|mm|abbr=on}} thick that is covered with sparse, bristled dark-brown to black hair. Short tactile hair grows on the trunk, which has two finger-like processes at the tip, whereas Asian elephants only have one.<ref name=Laurson1978/> Their large ears help to reduce body heat. Flapping them creates air currents and exposes the ears' inner sides where large blood vessels increase heat loss during hot weather. The trunk is a [[prehensile]] elongation of its upper lip and nose. This highly sensitive organ is innervated primarily by the [[trigeminal nerve]], and is thought to be manipulated by about 40,000–60,000 muscles. Because of this muscular structure, the trunk is so strong that elephants can use it to lift about 3% of their own body weight. They use it for smelling, touching, feeding, drinking, dusting, producing sounds, loading, defending and attacking.<ref name=Shoshani1978>{{cite journal |last1=Shoshani |first1=J. |year=1978 |title=General information on elephants with emphasis on tusks |journal=Elephant |volume=1 |issue=2 |pages=20–31 |doi=10.22237/elephant/1491234053 |url=https://digitalcommons.wayne.edu/cgi/viewcontent.cgi?article=1016&context=elephant|doi-access=free }}</ref> Elephants sometimes swim underwater and use their trunks as [[snorkel (swimming)|snorkels]].<ref name= "West2002">{{Cite journal|title= Why doesn't the elephant have a pleural space?|last= West|first= J.B.|date= April 2002|journal= News in Physiological Sciences |volume= 17|issue= 2|pages= 47–50|doi= 10.1152/nips.01374.2001|pmid= 11909991|s2cid= 27321751|doi-access= free}}</ref><ref>{{Cite web|last=Sandybinns|title=25 Things You Might Not Know About Elephants|url=https://elephantconservation.org/elephants/25-things-you-might-not-know-about-elephants/|access-date=2021-02-22|website=International Elephant Foundation|archive-date=27 September 2022|archive-url=https://web.archive.org/web/20220927060800/https://elephantconservation.org/elephants/25-things-you-might-not-know-about-elephants/|url-status=dead}}</ref>

=== Tusks and molars ===
Both male and female African elephants have tusks that grow from [[deciduous teeth]] called tushes, which are replaced by tusks when calves are about one year old. Tusks are composed of [[dentin]], which forms small diamond-shaped structures in the tusk's center that become larger at its periphery.<ref name=Shoshani1978/> Tusks are primarily used to dig for roots and strip the bark from trees for food, for fighting each other during the  mating season, and for defending themselves against predators. The tusks weigh from {{cvt|23|to(-)|45|kg|lb|0}} and can be from {{cvt|1.5|to(-)|2.4|m|ft|sigfig=1}} long. They are curved forward and continue to grow throughout the elephant's lifetime.<ref name=Burnie>{{cite book |last=Burnie |first=D. |title=Animal |year=2001 |publisher=[[Dorling Kindersley]] |location=London}}</ref>
[[File:Loxodonta africana - Molar of an adult.JPG|thumb|Molar of ''Loxodonta africana'', showing lozenge/loxodont]]
The [[dental formula]] of elephants is {{DentalFormula|upper=1.0.3.3|lower=0.0.3.3|total=26}}.<ref name="Shoshani1978" /> Elephants have four [[molars]]; each weighs about {{convert|5|kg|lb|abbr=on}} and measures about {{convert|30|cm|in|abbr=on}} long. As the front pair wears down and drops out in pieces, the back pair moves forward, and two new molars emerge in the back of the mouth. Elephants replace their teeth four to six times in their lifetimes. At around 40 to 60 years of age, the elephant loses the last of its molars and will likely die of starvation which is a common cause of death. African elephants have 24 teeth in total, six on each quadrant of the jaw. The [[Elephantidae|enamel plates]] of the molars are fewer in number than in Asian elephants.<ref name="Domestic">{{cite book |last=Clutton-Brock |first=Juliet |author-link=Juliet Clutton-Brock |title=A Natural History of Domesticated Mammals |year=1987 |isbn=0-521-34697-5 |page=208|publisher=Cambridge University Press }}</ref> The enamel of the molar teeth wears into a distinctive lozenge/loxodont ('''<>''') shape characteristic to all members of the genus ''Loxodonta''.''<ref name=":2" />'' While some extinct species of ''Loxodonta'' retained permanent [[premolar]] teeth, these have been lost in both living species.<ref>{{Cite journal |last=Sanders |first=William J. |date=2018-02-17 |title=Horizontal tooth displacement and premolar occurrence in elephants and other elephantiform proboscideans |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2017.1297436 |journal=Historical Biology |language=en |volume=30 |issue=1–2 |pages=137–156 |doi=10.1080/08912963.2017.1297436 |bibcode=2018HBio...30..137S |issn=0891-2963}}</ref>

=== Size ===
[[File:Jumbo size comparison.jpg|thumb|Skeleton of [[Jumbo]], a young African bush elephant bull, compared to a human]]
The African bush elephant is the largest [[terrestrial animal]]. Under optimal conditions where individuals are capable of reaching full growth potential, mature fully grown females are {{cvt|2.47–2.73|m}} tall at the shoulder and weigh {{cvt|2600–3500|kg|lb}}, while mature fully grown bulls are {{cvt|3.04–3.36|m}} tall and weigh {{cvt|5200–6900|kg|lb}} on average. The largest recorded bull stood {{cvt|3.96|m|ft|1}} at the shoulder and is estimated to have weighed {{cvt|10400|kg}}.<ref name=probos_mass>{{Cite journal |last1=Larramendi |first1=A. |year=2016 |title=Shoulder height, body mass and shape of proboscideans |journal=Acta Palaeontologica Polonica |volume=61 |issue=3 |pages=537–574 |doi=10.4202/app.00136.2014 |s2cid=2092950 |doi-access=free}}</ref>
Its back is concave-shaped, while the back of the African forest elephant is nearly straight.<ref name=Grubb_al2000/> The African forest elephant is considerably smaller. Fully grown African forest elephant males in optimal conditions where individuals are capable of reaching full growth potential are estimated to be on average {{Cvt|2.09–2.31|m|ft}} tall and {{Cvt|1700-2300|kg|lb}} in weight.<ref name=probos_mass/>

== Distribution and habitat ==
African elephants are distributed in [[Sub-Saharan Africa]], where they inhabit [[Sahel]]ian [[scrubland]] and [[arid]] regions, [[tropical rainforest]]s, [[mopane]] and [[miombo]] [[woodland]]s. African forest elephant populations occur only in Central and West Africa.<ref name="Blanc_al2003">{{cite book |last1=Blanc |first1=J. J. |last2=Thouless |first2=C. R. |last3=Hart |first3=J. A. |last4=Dublin |first4=H. T. |last5=Douglas-Hamilton |first5=I. |last6=Craig |first6=G. C. |last7=Barnes |first7=R. F. W. |title=African Elephant Status Report 2002: An update from the African Elephant Database |year=2003 |series=Occasional Paper of the IUCN Species Survival Commission No. 29 |publisher=IUCN |location=Gland and Cambridge |url=https://books.google.com/books?id=Z418bsp0fQoC|isbn=9782831707075 }}</ref>

== Behavior and ecology ==
=== Sleeping pattern===
Elephants are the animals with the lowest sleep times, especially African elephants. Research has found their average sleep to be only 2 hours in 24-hour cycles.

=== Family ===
[[File:Serengeti Elefantenherde2.jpg|thumb|Female bush elephants in [[Tanzania]]]]
[[File:Bee-Threat-Elicits-Alarm-Call-in-African-Elephants-pone.0010346.s004.ogv|thumb|Family responds to bee warning rumble]]
Both African elephant species live in family units comprising several adult cows, their daughters and their subadult sons. Each family unit is led by an older cow known as the [[matriarch]].<ref name=Douglas-Hamilton1972>{{cite thesis |last1=Douglas-Hamilton |first1=I. |year=1972 |title=On the ecology and behaviour of the African elephant: the elephants of Lake Manyara |type=PhD thesis |publisher=University of Oxford |location=Oxford |id={{EThOS|uk.bl.ethos.453899}} }}</ref><ref name=Turkalo_al2013>{{cite book |editor=Kingdon, J. |editor2=Happold, D. |editor3=Hoffmann, M. |editor4=Butynski, T. |editor5=Happold, M. |editor6=Kalina, J. |year=2013 |title=The Mammals of Africa |volume=I. Introductory Chapters and Afrotheria |publisher=Bloomsbury Publishing |location=London |author=Turkalo, A.|author2=Barnes, R. |chapter=''Loxodonta cyclotis'' Forest Elephant |pages=195–200
|isbn=9781408189962 |chapter-url=https://books.google.com/books?id=B_07noCPc4kC&pg=PA195}}</ref> African forest elephant groups are less cohesive than African bush elephant groups, probably because of the lack of predators.<ref name=Turkalo_al2013/>

When separate family units bond, they form kinship or bond groups. After [[puberty]], male elephants tend to form close alliances with other males. While females are the most active members of African elephant groups, both male and female elephants are capable of distinguishing between hundreds of different low-frequency [[infrasonic]] calls to communicate with and identify each other.<ref>{{Cite journal |last1=McComb |first1=K. |last2=Moss |first2=C. |last3=Sayialel|first3=S. |last4=Baker|first4=L. |date=2000 |title=Unusually extensive networks of vocal recognition in African elephants |journal=Animal Behaviour |volume=59 |issue=6|pages=1103–1109 |doi=10.1006/anbe.2000.1406|pmid=10877888|s2cid=17628123 |citeseerx=10.1.1.332.7064 }}</ref><ref>{{Cite journal |last1=McComb |first1=K. |last2=Moss|first2=C.|last3=Durant|first3=S. M. |last4=Baker |first4=L. |last5=Sayialel |first5=S. |date=2001 |title=Matriarchs As Repositories of Social Knowledge in African Elephants |journal=Science |volume=292 |issue=5516 |pages=491–494 |doi=10.1126/science.1057895|pmid=11313492 |bibcode=2001Sci...292..491M|s2cid=17594461 }}</ref>

Elephants use [[Infrasound#Animal reactions to infrasound|some vocalisations]] that are beyond the hearing range of humans,<ref>{{cite journal | last1 = Herbst | first1 = C. T. | last2 = Stoeger | first2 = A. S. | last3 = Frey | first3 = R. | last4 = Lohscheller | first4 = J. | last5 = Titze | first5 = I. | last6 = Gumpenberger | first6 = M. | last7 = Fitch | first7 = W. T. |year = 2012 | title = How Low Can You Go? Physical Production Mechanism of Elephant Infrasonic Vocalizations | doi =  10.1126/science.1219712  | journal = Science | volume = 337 | issue = 6094| pages = 595–599 | pmid=22859490| bibcode = 2012Sci...337..595H| s2cid = 32792564 }}</ref> to communicate across large distances. Elephant mating rituals include the gentle entwining of trunks.<ref>{{cite book |last1=Buss |first1=I. |last2=Smith |first2=N. |title=Observations on Reproduction and Breeding Behavior of the African Elephant |publisher=Allen Press |year=1966 |url=http://faculty.rio.edu/dalthoff/Courses/MA/TP-Buss%20and%20Smith-Elephant%20Repo%20Breeding%20Behavior-1966-JWM.pdf |access-date=8 March 2013 |archive-url=https://web.archive.org/web/20150518072553/http://faculty.rio.edu/dalthoff/Courses/MA/TP-Buss%20and%20Smith-Elephant%20Repo%20Breeding%20Behavior-1966-JWM.pdf |archive-date=18 May 2015 |url-status=dead}}</ref>

Bulls were believed to be solitary animals, becoming independent once reaching maturity. New research suggests that bulls maintain ecological knowledge for the herd, facilitating survival when searching for food and water, which also benefits the young bulls who associate with them. Bulls only return to the herd to breed or to socialize; they do not provide prenatal care to their offspring, but rather play a fatherly role to younger bulls to show dominance.<ref>{{Cite news|last=Briggs|first=H. |date=2020 |title=Secrets of male elephant society revealed in the wild |work=BBC News|url=https://www.bbc.com/news/science-environment-54018133|access-date=2020-09-07}}</ref>

=== Feeding ===
While feeding, the African elephant uses its trunk to pluck leaves and its tusks to tear at branches, which can cause enormous damage to foliage.<ref name=Burnie /> [[Fermentation]] of the food takes place in the [[hindgut]], enabling large food intakes.<ref name=Clauss>{{cite journal |last1=Clauss |first1=M. |last2=Frey |first2=R. |last3=Kiefer |first3=B. |last4=Lechner-Doll |first4=M. |last5=Loehlein |first5=W. |last6=Polster |first6=C. |last7=Roessner |first7=G. E. |last8=Streich |first8=W. J. |title=The maximum attainable body size of herbivorous mammals: morphophysiological constraints on foregut, and adaptations of hindgut fermenters |journal=[[Oecologia]] |volume=136 |issue=1 |pages=14–27 |date=2003 |doi=10.1007/s00442-003-1254-z |pmid=12712314 |bibcode=2003Oecol.136...14C |s2cid=206989975 |url=http://www.zora.uzh.ch/id/eprint/2393/2/Oecologia_body_size_2003V.pdf |access-date=24 September 2019 |archive-date=8 June 2019 |archive-url=https://web.archive.org/web/20190608152406/https://www.zora.uzh.ch/id/eprint/2393/2/Oecologia_body_size_2003V.pdf |url-status=dead }}</ref> The large size and hindgut of the African elephant also allows for the digestion of various plant parts, including fibrous stems, bark and roots.<ref>{{Cite journal|last1=Owen-Smith|first1=Norman|last2=Chafota|first2=Jonas|date=2012-06-28|title=Selective feeding by a megaherbivore, the African elephant (''Loxodonta africana'')|journal=Journal of Mammalogy|volume=93|issue=3|pages=698–705|doi=10.1644/11-MAMM-A-350.1|s2cid=83726553|doi-access=free}}</ref>

=== Intelligence ===
{{See also|Elephant cognition}}
[[File:Elephant scratching on a tree.jpg|thumb|Scratching on a tree helps to remove layers of dead skin and [[Parasitism|parasites]]]]
African elephants are highly intelligent.<ref>{{cite journal |author1=Plotnik, J. M. |author2=De Waal, F. B. |author3=Reiss, D. |date=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 |pmc=1636577 |bibcode=2006PNAS..10317053P |doi-access=free }}</ref> They have a very large and highly convoluted [[neocortex]], a trait they share with [[humans]], [[apes]] and some [[dolphin]] species. They are among the world's most intelligent species. With a mass of just over {{cvt|5|kg|abbr=on}}, the [[Elephant cognition#Brain structure|elephant brain]] is larger than that of any other terrestrial animal. The elephant's brain is similar to a [[human brain]] in terms of structure and complexity; the elephant's [[cerebral cortex|cortex]] has as many [[neurons]] as that of a human brain,<ref name=Roth>{{cite book |last1=Roth |first1=G. |last2=Stamenov |first2=M. I. |last3=Gallese |first3=V. |chapter=Is the human brain unique? |title=Mirror Neurons and the Evolution of Brain and Language |publisher=John Benjamins Publishing |pages=63–76 |doi=10.1002/0470867221.ch2 |isbn=978-0-470-84960-6|year=2003 }}</ref> suggesting [[convergent evolution]].<ref>{{cite journal |last1=Goodman |first1=M. |last2=Sterner |first2=K. |last3=Islam |first3=M. |last4=Uddin |first4=M. |last5=Sherwood |first5=C. |last6=Hof |first6=P. |last7=Hou |first7=Z. |last8=Lipovich |first8=L. |last9=Jia |first9=H. |last10=Grossman |first10=L. |last11=Wildman |first11=D. |title=Phylogenomic analyses reveal convergent patterns of adaptive evolution in elephant and human ancestries |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=49 |pages=20824–20829 |date=2009 |doi=10.1073/pnas.0911239106 |pmc=2791620 |pmid=19926857 |bibcode=2009PNAS..10620824G |doi-access=free}}</ref>

Elephants exhibit a wide variety of behaviours, including those associated with [[grief]], [[learning]], [[mimicry]], [[art]], [[Play (activity)|play]], a sense of [[humor]], [[altruism]], use of [[tools]], [[compassion]], [[cooperation]],<ref name=PNAS>{{cite journal |title=Elephants know when they need assistance in a cooperative task |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=12 |pages=5116–5121 |doi=10.1073/pnas.1101765108 |pmid=21383191 |pmc=3064331 |date=2011 |last2=Suphachoksahakun |first2=W. |last3=Lair |first3=R. |last4=Plotnik |first4=J. M. |last1=Plotnik |first1=J. M. |doi-access=free}}</ref> [[self-awareness]], [[memory]] and possibly [[language]].<ref name=NatlGeoNews>{{cite news |last=Parsell |first=D. L. |title=In Africa, Decoding the 'Language' of Elephants |publisher=[[National Geographic Magazine]] News |date=2003 |url=http://news.nationalgeographic.com/news/2003/02/0221_030221_elephantvocal1.html |access-date=30 October 2007 |archive-url=https://web.archive.org/web/20071012170607/http://news.nationalgeographic.com/news/2003/02/0221_030221_elephantvocal1.html |archive-date=12 October 2007 |url-status=dead}}</ref> All of these behaviors point to a highly intelligent species that is thought to be equal to [[cetaceans]]<ref name=ABCScience>{{cite web |last=Viegas |first=J. |title=Elephants smart as chimps, dolphins |work=ABC Science |year=2011 |access-date=8 March 2011 |url=http://www.abc.net.au/science/articles/2011/03/08/3158077.htm}}</ref><ref name="Jennifer Viegas 2011">{{cite web |last=Viegas |first=J. |title=Elephants Outwit Humans During Intelligence Test |work=Discovery News |year=2011 |url=http://news.discovery.com/animals/elephants-intelligence-test-110307.html |access-date=19 March 2011 |archive-url=https://web.archive.org/web/20110308093715/http://news.discovery.com/animals/elephants-intelligence-test-110307.html |archive-date=8 March 2011 |url-status=live}}</ref> and [[primate]]s.<ref name="Jennifer Viegas 2011" />

The African elephant’s cognitive complexity includes behaviors indicative of empathy, problem-solving, and cooperative group behaviors. These traits underscore the evolutionary convergence of intelligence across species, similar to that seen in primates and cetaceans.<ref>{{cite journal |last1=Plotnik |first1=J. M. |last2=Suphachoksahakun |first2=W. |title=Elephants know when they need assistance in a cooperative task |journal=Proceedings of the National Academy of Sciences |volume=108 |issue=12 |pages=5116–5121 |doi=10.1073/pnas.1101765108 |pmid=21383191 |pmc=3064331 |date=2011|doi-access=free }}</ref>

=== Reproduction ===
{{multiple image|perrow=1|image1=Bull elephants small.jpg|caption1=Bull elephants in mock aggression|image2=Elephant (Loxodonta africana) mating ritual composite.jpg|caption2=[[Courtship display|Mating ritual]]}}
African elephants are at their most [[fertility|fertile]] between the ages of 25 and 45. Calves are born after a [[gestation]] period of up to nearly two years.<ref name="Macdonald">{{cite book |last=Macdonald |first=D. |title=The New Encyclopedia of Mammals |publisher=[[Oxford University Press]] |year=2001 |location=Oxford |chapter=}}</ref> The calves are cared for by their mother and other young females in the group, known as [[allomothering]].<ref name=Douglas-Hamilton1972/>

African elephants show [[sexual dimorphism]] in weight and shoulder height by age 20, due to the rapid early growth of males. By age 25, males are double the weight of females; however, both sexes continue to grow throughout their lives.

Female African elephants are able to start reproducing at around 10 to 12 years of age,<ref>{{cite journal |last=Benedict |first=F. G. |title=The physiology of the elephant |year=1936 |journal=Carnegie Inst. Washington Pub. No. 474 |volume=1}}</ref> and are in estrus for about 2 to 7 days. They do not mate at a specific time; however, they are less likely to reproduce in times of drought than when water is plentiful. The gestation period of an elephant is 22 months and fertile females usually give birth every 3–6 years, so if they live to around 50 years of age, they may produce 7 offspring. Females are a scarce and mobile resource for the males so there is intense competition to gain access to estrous females.

Post sexual maturity, males begin to experience [[musth]], a physical and behavioral condition that is characterized by elevated testosterone, aggression and more sexual activity.<ref name=Hollister-Smith>{{cite journal|last1=Hollister-Smith |first1=J. A. |last2=Poole |first2=J. H. |last3=Archie |first3=E. A. |last4=Vance |first4=E. A. |last5=Georgiadis |first5=N. J. |last6=Moss |first6=C. J. |author-link6=Cynthia Moss |last7=Alberts |first7=S. C. |title=Age, musth, and paternity success in wild male African elephants, ''Loxodonta africana'' |journal=Animal Behaviour |date=2007 |volume=74 |issue=2 |pages=287–296 |doi=10.1016/j.anbehav.2006.12.008 |s2cid=54327948 |url=http://sites.biology.duke.edu/albertslab/pdfs/Hollister-Smith%26al_2007.pdf|archive-url=https://web.archive.org/web/20150518085653/http://sites.biology.duke.edu/albertslab/pdfs/Hollister-Smith%26al_2007.pdf |archive-date=18 May 2015 }}</ref><ref name=Sukumar2003>{{cite book |last=Sukumar |first=R. |author-link=Raman Sukumar |title=The Living Elephants: Evolutionary Ecology, Behavior, and Conservation |publisher=Oxford University Press |location=Oxford, USA |date=2003 |isbn=0-19-510778-0 |pages=112–124 |chapter=Sexual selection and mate choice}}</ref> Musth also serves a purpose of calling attention to the females that they are of good quality, and it cannot be mimicked as certain calls or noises may be. Males sire few offspring in periods when they are not in musth. During the middle of estrus, female elephants look for males in musth to guard them. The females will yell, in a loud, low way to attract males from far away. Male elephants can also smell the hormones of a female ready for breeding. This leads males to compete with each other to mate, which results in the females mating with older, healthier males.<ref name = Poole1989>{{cite journal |last=Poole |first=Joyce H. |title=Mate guarding, reproductive success and female choice in African elephants |journal=Animal Behaviour |year=1989 |volume=37 |pages=842–849 |doi=10.1016/0003-3472(89)90068-7|s2cid=53150105 }}</ref> Females choose to a point who they mate with, since they are the ones who try to get males to compete to guard them. However, females are not guarded in the early and late stages of estrus, which may permit mating by younger males not in musth.<ref name = Poole1989/>

Males over the age of 25 compete strongly for females in estrus, and are more successful the larger and more aggressive they are.<ref name=Sukumar2003/> Bigger males tend to sire bigger offspring.<ref>{{cite journal |last1=Lee |first1=Phyllis C. |last2=Moss |first2=Cynthia J. |title=Early maternal investment in male and female African elephant calves |journal=Behavioral Ecology and Sociobiology |year=1986 |volume=18 |issue=5 |pages=353–361 |doi=10.1007/bf00299666|bibcode=1986BEcoS..18..353L |s2cid=10901693}}</ref> Wild males begin breeding in their thirties when they are at a size and weight that is competitive with other adult males. Male reproductive success is maximal in mid-adulthood and then begins to decline. However, this can depend on the ranking of the male within their group, as higher-ranking males maintain a higher rate of reproduction.<ref>{{cite journal |last1=Loizi |first1=H. |last2=Goodwin |first2=T. E. |last3=Rasmussen |first3=L. E. L. |last4=Whitehouse |first4=A. M. |last5=Schulte |first5=B. A. |title=Sexual dimorphism in the performance of chemosensory investigatory behaviours by African elephants (''Loxodonta africana'') |year=2009 |journal=Behaviour |volume=146 |issue=3 |pages=373–392 |doi=10.1163/156853909X410964}}</ref> Most observed matings are by males in musth over 35 years of age. Twenty-two long observations showed that age and musth are extremely important factors; "… older males had markedly elevated paternity success compared with younger males, suggesting the possibility of sexual selection for longevity in this species."<ref name=Hollister-Smith/>{{rp|287}}

Males usually stay with a female and her herd for about a month before moving on in search of another mate. Less than a third of the population of female elephants will be in estrus at any given time and the gestation period of an elephant is long, so it makes more evolutionary sense for a male to search for as many females as possible rather than stay with one group.{{citation needed|date=April 2019}}

== Threats ==
[[File:Number of African elephants.svg|thumb|Number of African elephants]]
[[File:Ivory trade.jpg|thumb|upright|Men with African elephant tusks in [[Dar es Salaam]], {{Circa|1900|lk=yes}}]]
Both species are threatened by [[habitat loss]] and fragmentation, and [[poaching]] for the illegal [[ivory trade]] is a threat in several range countries as well. The African bush elephant is listed as [[Endangered species|Endangered]] and the African forest elephant as [[Critically Endangered]] on the respective [[IUCN Red List]]s.<ref>{{cite iucn |author=Gobush, K.S. |author2=Edwards, C.T.T. |author3= Balfour, D. |author4=Wittemyer, G. |author5=Maisels, F. |author6=Taylor, R.D. |year=2022 |amends=2021 |title=''Loxodonta africana'' |page=e.T181008073A223031019 |doi=10.2305/IUCN.UK.2022-2.RLTS.T181008073A223031019.en}}</ref><ref>{{cite iucn |author=Gobush, K.S. |author2=Edwards, C.T.T. |author3= Maisels, F. |author4=Wittemyer, G. |author5=Balfour, D. |author6=Taylor, R.D. |year=2021 |errata=2021 |title=''Loxodonta cyclotis'' |page=e.T181007989A204404464 |doi=10.2305/IUCN.UK.2021-1.RLTS.T181007989A204404464.en}}</ref>

Based on vegetation types that provide suitable habitat for African elephants, it was estimated that in the early 19th century a maximum of 26,913,000 African elephants might have been present from the [[Sahel]] in the north to the [[Highveld]] in the south. Decrease of suitable habitat was the major cause for the decline of elephant populations until the 1950s. Hunting African elephants for the [[ivory trade]] accelerated the decline from the 1970s onwards. The [[carrying capacity]] of remaining suitable habitats was estimated at 8,985,000 elephants at most by 1987.<ref>{{cite journal |last1=Milner-Gulland |first1=E. J. |last2=Beddington |first2=J. R. |year=1993 |title=The exploitation of elephants for the ivory trade: an historical perspective |journal=Proceedings of the Royal Society B: Biological Sciences |volume=252 |issue=1333 |pages=29–37 |doi=10.1098/rspb.1993.0042 |url=https://www.researchgate.net/publication/252970898 |bibcode=1993RSPSB.252...29M |s2cid=128704128}}</ref>
In the 1970s and 1980s, the price of ivory rose, and poaching for ivory increased,  particularly in Central African range countries where access to elephant habitats was facilitated by [[logging]] and [[petroleum]] mining industries.<ref name=Blanc_al2003 />
Between 1976 and 1980, about {{convert|830|t|abbr=on}} raw ivory was exported from Africa to [[Hong Kong]] and [[Japan]], equivalent to tusks of about 222,000 African elephants.<ref>{{cite journal |last1=Parker |first1=J. S. C. |last2=Martin |first2=E. B. |year=1982 |title=How many elephants are killed for the ivory trade? |journal=Oryx |volume=16 |issue=3 |pages=235–239 |url=http://www.rhinoresourcecenter.com/pdf_files/133/1337589439.pdf |doi=10.1017/S0030605300017452 |doi-access=free}}</ref>

The first continental elephant census was carried out in 1976. At the time, 1.34 million elephants were estimated to range over {{cvt|7300000|sqkm}}.<ref name=Stiles2004>{{cite journal |last1=Stiles |first1=D. |year=2004 |title=The ivory trade and elephant conservation |journal=Environmental Conservation |volume=31 |issue=4 |pages=309–321 |doi=10.1017/S0376892904001614 |bibcode=2004EnvCo..31..309S |s2cid=14772884 |url=http://danstiles.org/publications/ivory/09.Env.Cons.final.pdf}}</ref>
In the 1980s, it was difficult to carry out systematic surveys in several East African range countries due to [[civil war]]s.<ref name=Blanc_al2003 />
In 1987, it was estimated that the African elephant population had declined to 760,000 individuals. In 1989, only 608,000 African elephants were estimated to have survived.<ref name=Stiles2004/>
In 1989, the Kenyan Wildlife Service burned a stockpile of tusks in protest against the ivory trade.<ref>{{cite book |last=Poole |first=J. |title=Coming of Age With Elephants |year=1996 |publisher=Hyperion |location=New York |isbn=0-7868-6095-2 |page=[https://archive.org/details/isbn_9780786860951/page/232 232] |url=https://archive.org/details/isbn_9780786860951/page/232}}</ref>

When the international ivory trade reopened in 2006, the demand and price for ivory increased in Asia. In Chad's [[Zakouma National Park]], more than 3,200 elephants were killed between 2005 and 2010. The park did not have sufficient guards to combat poaching and their weapons were outdated. Well organized networks facilitated smuggling the ivory through Sudan.<ref name=Poilecot2010>{{cite journal |last1=Poilecot |first1=P. |year=2010 |title= Le braconnage et la population d'éléphants au Parc National de Zakouma (Tchad) |journal=Bois et Forêts des Tropiques |volume=303 |issue=303 |pages=93–102 |doi=10.19182/bft2010.303.a20454 |url=https://www.researchgate.net/publication/291781119 |doi-access=free}}</ref>
The government of Tanzania estimated that more than 85,000 elephants were lost to poaching in Tanzania between 2009 and 2014, representing a 60% loss.<ref name=eal>{{cite news |last=Mathiesen |first=K. |date=2015 |title=Tanzania elephant population declined by 60% in five years, census reveals |url=https://www.theguardian.com/environment/2015/jun/02/tanzania-epicentre-of-elephant-poaching-census-reveals |work=[[The Guardian]] |location=London |access-date= 23 August 2017}}</ref>
In 2012, a large upsurge in ivory poaching was reported, with about 70% of the product flowing to China.<ref name=nyt120903>{{cite news |last=Gettleman |first=J. |author-link=Jeffrey Gettleman |title=Elephants Dying in Epic Frenzy as Ivory Fuels Wars and Profits |work=The New York Times |date=3 September 2012 |url=https://www.nytimes.com/2012/09/04/world/africa/africas-elephants-are-being-slaughtered-in-poaching-frenzy.html}}</ref>
China was the biggest market for poached ivory but announced that it would phase out the legal domestic manufacture and sale of ivory products in May 2015.<ref>{{cite news |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|author=Fergus Ryan|newspaper=The Guardian|date=26 September 2015}}</ref>

Conflicts between elephants and a growing human population are a major issue in elephant conservation.<ref name=Blanc_al2003 /> Human encroachment into natural areas where bush elephants occur or their increasing presence in adjacent areas has spurred research into methods of safely driving groups of elephants away from humans. Playback of the recorded sounds of angry [[Western honey bee]]s has been found to be remarkably effective at prompting elephants to flee an area.<ref>{{cite journal |last1=King |first1=L. E. |last2=Douglas-Hamilton |first2=I. |author-link2=Iain Douglas-Hamilton |last3=Vollrath |first3=F. |title=African elephants run from the sound of disturbed bees |year=2007 |journal=Current Biology |volume=17 |issue=19 |pages=R832–R833 |doi=10.1016/j.cub.2007.07.038 |pmid=17925207|s2cid=30014793 |doi-access=free|bibcode=2007CBio...17.R832K }}</ref> Farmers have tried scaring elephants away by more aggressive means such as fire or the use of chili peppers along fences to protect their crops.<ref>{{Cite journal |last1=Snyder|first1=K. |last2=Mneney |first2=P. |last3=Benjamin |first3=B. |last4=Mkilindi|first4=P. |last5=Mbise|first5=N. |date=2019|title=Seasonal and spatial vulnerability to agricultural damage by elephants in the western Serengeti, Tanzania |journal=Oryx |volume=55 |issue=1 |pages=139–149 |doi=10.1017/S0030605318001382 |doi-access=free |url=https://www.cambridge.org/core/services/aop-cambridge-core/content/view/25D9D4A887EEB3918933EE5AA361478D/S0030605318001382a_hi.pdf/seasonal-and-spatial-vulnerability-to-agricultural-damage-by-elephants-in-the-western-serengeti-tanzania.pdf}}</ref>

== Conservation ==
In 1986, the African Elephant Database was initiated with the aim to monitor the status of African elephant populations. This database includes results from aerial surveys, dung counts, interviews with local people and data on poaching.<ref name=AfESG2016>{{cite book |last1=Thouless |first1=C. R. |last2=Dublin |first2=H. T. |last3=Blanc |first3=J. J. |last4=Skinner |first4=D. P. |last5=Daniel |first5=T. E. |last6=Taylor |first6=R. D. |last7=Maisels |first7=F. |last8=Frederick |first8=H. L. |last9=Bouché |first9=P. |year=2016 |title=African Elephant Status Report 2016 : an update from the African Elephant Database |publisher=IUCN SSC African Elephant Specialist Group |location=Gland |isbn=978-2-8317-1813-2 |series=Occasional Paper of the IUCN Species Survival Commission No. 60 |url=https://www.dropbox.com/s/7a8w3kk6r9hzm0r/AfESG%20African%20Elephant%20Status%20Report%202016.pdf?dl=1 }}{{Dead link|date=August 2023 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>

In 1989, the [[Convention on International Trade in Endangered Species of Wild Fauna and Flora]] listed the African elephant on [[CITES Appendix I]]. This listing banned international trade of African elephants and their body parts by countries that signed the CITES agreement. Hunting elephants is banned in the Central African Republic, Democratic Republic of Congo, Gabon, Côte d'Ivoire, and Senegal. After the ban came into force in 1990, retail sales of [[ivory carving]]s in South Africa have plummeted by more than 95% within 10 years.<ref>{{cite journal |last1=Stiles |first1=D. |last2=Martin |first2=E. |year=2001 |title=Status and trends of the ivory trade in Africa, 1989–1999 |journal=Pachyderm |issue=30 |pages=24–36 |url=https://www.iucn.org/backup_iucn/cmsdata.iucn.org/downloads/pachy30.pdf#page=27 |access-date=8 June 2019 |archive-date=3 January 2020 |archive-url=https://web.archive.org/web/20200103093454/https://www.iucn.org/backup_iucn/cmsdata.iucn.org/downloads/pachy30.pdf#page=27 |url-status=dead }}</ref>
As a result of the trade ban, African elephant populations recovered in [[Southern Africa]]n range countries.<ref>{{cite book |last1=Blanc  |first1=J. J. |last2=Barnes |first2=R. F. W. |last3=Craig |first3=G. C. |last4=Dublin |first4=H. T. |last5=Thouless |first5=C. R. |last6=Douglas-Hamilton |first6=I. |last7=Hart |first7=J. A. |year=2007 |title=African Elephant Status Report 2007: An update from the African Elephant Database |publisher=IUCN SSC African Elephant Specialist Group |location=Gland |series=Occasional Paper of the IUCN Species Survival Commission No. 33 |url=https://portals.iucn.org/library/efiles/documents/SSC-OP-033.pdf}}</ref>

The African Elephant Specialist Group has set up a ''Human-Elephant Conflict Task Force'' with the aim to develop conflict mitigation strategies.<ref>{{cite book |author1=Naughton, L. |author2=Rose, R. |author3=Treves, A. |title=The social dimensions of human-elephant conflict in Africa: A literature review and case studies from Uganda and Cameroon |year=1999 |publisher=University of Wisconsin |location=Madison |url=https://www.researchgate.net/publication/228594746}}</ref>

In 2005, the [[West African Elephant Memorandum of Understanding]] was signed by 12 West African countries. The [[Convention on the Conservation of Migratory Species of Wild Animals]] provided financial support for four years to implement the West African Elephant Conservation Strategy, which forms the central component of this intergovernmental treaty.<ref>{{cite journal |last1=Dublin |first1=H. |year=2005 |title=African Elephant Specialist Group report |journal=Pachyderm |issue=39 |pages=1–9 |url=https://library.wur.nl/WebQuery/wurpubs/fulltext/17713}}</ref>
[[File:Elephant african mask.jpg|thumb|right|Elephant mask in the [[Ivory Coast]]]]
In 2019, the export of wild African elephants to zoos around the world was banned, with an exception added by the EU to allow export in "exceptional cases where … it is considered that a transfer to ex-situ locations will provide demonstrable in-situ conservation benefits for African elephants". Previously, export had been allowed in Southern Africa with Zimbabwe capturing and exporting more than 100 baby elephants to Chinese zoos since 2012.<ref>{{cite news |title=Near-total ban imposed on sending wild African elephants to zoos |url=https://www.theguardian.com/environment/2019/aug/27/near-total-ban-imposed-on-sending-wild-african-elephants-to-zoos}}</ref>

It was found that elephant conservation does not pose a trade-off with [[climate change mitigation]]. Although animals typically cause a reduction of woody biomass and therewith above-ground carbon, they foster soil [[carbon sequestration]].<ref>{{Cite journal|last1=Sandhage-Hofmann |first1=A. |last2=Linstädter |first2=A. |last3=Kindermann |first3=L. |last4=Angombe |first4=S. |last5=Amelung |first5=W. |date=2021|title=Conservation with elevated elephant densities sequesters carbon in soils despite losses of woody biomass |journal=Global Change Biology |volume=27 |issue=19 |pages=4601–4614 |doi=10.1111/gcb.15779|pmid=34197679 |doi-access=free }}</ref>

==In culture==
Many [[Culture of Africa|African cultures]] revere the African elephant as a symbol of strength and power.<ref>{{cite web|title=383. African Elephant (''Loxodonta africana'') |url=http://www.edgeofexistence.org/mammals/species_info.php?id=77 |publisher=EDGE: Mammal Species Information|access-date=7 December 2012}}</ref><ref>{{cite web|title=West African Elephants|url=http://www.cms.int/species/elephants/index.htm|publisher=Convention on Migratory Species|access-date=7 December 2012}}</ref> It is also praised for its size, longevity, stamina, mental faculties, cooperative spirit, and [[loyalty]].<ref>{{cite web|title=Elephant: The Animal and Its Ivory in African Culture |url=http://www.fowler.ucla.edu/category/exhibitions-education/elephant-animal-and-its-ivory-african-culture |publisher=Fowler Museum at UCLA |access-date=24 January 2013 |url-status=dead |archive-url=https://web.archive.org/web/20130330072035/http://www.fowler.ucla.edu/category/exhibitions-education/elephant-animal-and-its-ivory-african-culture |archive-date=30 March 2013 }}</ref> Its religious importance is mostly [[Totemism|totemic]].<ref>{{cite book|author=Sukumar, R. |date=2003 |title=The Living Elephants: Evolutionary Ecology, Behaviour, and Conservation |chapter=The capture and use of the African elephant |pages=81–88 |publisher=Oxford University Press |place=Oxford, USA |isbn=978-0-19-510778-4 |oclc= 935260783}}</ref> Many societies believed that their chiefs would be [[reincarnation|reincarnated]] as elephants. In the 10th century, the people of [[Igbo-Ukwu]] in Nigeria buried their leaders with elephant tusks.<ref>{{cite book|author= Wylie, D.|title= Elephant |url=https://books.google.com/books?id=Aj75snWdfv8C |page=79 |date=2009|publisher= Reaktion Books|isbn= 978-1-86189-615-5 |oclc= 740873839}}</ref>

[[South Africa]] uses elephant tusks in their [[Coat of arms of South Africa|coat of arms]] to represent wisdom, strength, moderation and eternity.<ref>{{cite web|title=National Coat of Arms |url=http://www.info.gov.za/aboutgovt/symbols/coa/index.htm |publisher=South African Government Information |access-date=7 December 2012 |url-status=dead |archive-url=https://web.archive.org/web/20120904094036/http://www.info.gov.za/aboutgovt/symbols/coa/index.htm |archive-date=4 September 2012}}</ref>

In the western African [[Kingdom of Dahomey]], the elephant was associated with the 19th century rulers of the [[Fon people]], [[Guezo]] and his son [[Glele]].{{efn|Guezo and Glele ruled from 1818 to 1858 and from 1858 to 1889, respectively}} The animal is believed to evoke strength, royal legacy, and enduring memory as related by the proverbs: "''There where the elephant passes in the forest, one knows''" and "''The animal steps on the ground, but the elephant steps down with strength''."<ref>{{cite web|title=Elephant Figure {{!}} Fon peoples {{!}} The Met|url=https://www.metmuseum.org/art/collection/search/318417|website=The Metropolitan Museum of Art|access-date=12 December 2017}}</ref> Their flag depicted an elephant wearing a royal crown.

===As national symbols===
The [[coat of arms of the Central African Republic]] features the head of an elephant in the upper left quadrant of the shield. The version of the [[coat of arms of Guinea]] used from 1958 to 1984 featured a golden elephant in the centre of the shield. The [[coat of arms of Ivory Coast]] features the head of an elephant as the focal point of the emblem. The [[coat of arms of the Republic of the Congo]] has two elephants supporting the shield. The [[coat of arms of Eswatini]] has an elephant and a lion supporting the shield.

==See also==
*''[[Africa's Elephant Kingdom]]''
*[[Indian elephant]]
*[[List of individual elephants]]
*[[Sri Lankan elephant]]
*[[Sumatran elephant]]

==Notes==
{{reflist|group=lower-alpha}}

== References ==
{{Reflist}}

== External links ==
*{{cite web |url=https://www.cms.int/en/species/loxodonta-africana |title=''Loxodonta africana'' |website=Convention on the Conservation of Migratory Species of Wild Animals |date=2020}} 
* [http://elephant.elehost.com/About_Elephants/about_elephants.htm Elephant Information Repository] {{Webarchive|url=https://web.archive.org/web/20090318112402/http://elephant.elehost.com/About_Elephants/about_elephants.htm |date=18 March 2009 }} – An in-depth resource on elephants
* [http://faculty.jsd.claremont.edu/dmcfarlane/MtElgon/index.htm "Elephant caves" of Mt Elgon National Park]
* [https://elephantvoices.org/ ElephantVoices] – Resource on elephant vocal communications
* [https://www.elephanttrust.org/ Amboseli Trust for Elephants] – Interactive web site
* [https://web.archive.org/web/20151107212924/http://anotherelephant.org/ Another Elephant] – A hub for saving the elephants.
*{{cite magazine |author=David Quammen |title=Family ties – The elephants of Samburu |magazine=National Geographic Magazine |date=2008 |url=http://ngm.nationalgeographic.com/2008/09/samburu-elephants/quammen-text|archive-url=https://web.archive.org/web/20080826010035/http://ngm.nationalgeographic.com/2008/09/samburu-elephants/quammen-text|archive-date=August 26, 2008}}
* [https://eia-international.org/ EIA 25 yrs investigating the ivory trade, reports etc]
* [https://us.eia.org/ EIA (in the USA) reports etc]
* [https://elephantconservation.org/elephants/african-elephants/ International Elephant Foundation]
* [https://www.worldwildlife.org/species/african-elephant]
* [https://pubmed.ncbi.nlm.nih.gov/28249035/]

{{Proboscidea|state=expanded}}
{{Proboscidea Genera}}
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[[Category:African elephants| ]]
[[Category:Mammals of Sub-Saharan Africa]]
[[Category:EDGE species]]
[[Category:Animal species groups]]
[[Category:National symbols of the Central African Republic]]
[[Category:National symbols of Equatorial Guinea]]
[[Category:National symbols of Ghana]]
[[Category:National symbols of Guinea]]
[[Category:National symbols of Ivory Coast]]
[[Category:National symbols of the Republic of the Congo]]
[[Category:National symbols of São Tomé and Príncipe]]
[[Category:National symbols of Eswatini]]
[[Category:Articles containing video clips]]
[[Category:Extant Pliocene first appearances]]