Elephantidae

{{Hatnote|For detailed information on the anatomy of living elephants, see Elephant#Anatomy}}

Elephantids are distinguished from more basal proboscideans like gomphotheres by their teeth, which have parallel lophs, formed from the merger of the cusps found in the teeth of more basal proboscideans, which are bound by cementum.{{Cite journal |last=Lister |first=Adrian M. |date=2013-06-26 |title=The role of behaviour in adaptive morphological evolution of African proboscideans |url=http://dx.doi.org/10.1038/nature12275 |journal=Nature |volume=500 |issue=7462 |pages=331–334 |doi=10.1038/nature12275 |pmid=23803767 |bibcode=2013Natur.500..331L |s2cid=883007 |issn=0028-0836|url-access=subscription }} In later elephantids, these lophs became narrow lamellae/plates,{{Cite journal |last1=Saarinen |first1=Juha |last2=Lister |first2=Adrian M. |date=2023-08-14 |title=Fluctuating climate and dietary innovation drove ratcheted evolution of proboscidean dental traits |journal=Nature Ecology & Evolution |language=en |volume=7 |issue=9 |pages=1490–1502 |bibcode=2023NatEE...7.1490S |doi=10.1038/s41559-023-02151-4 |issn=2397-334X |pmc=10482678 |pmid=37580434}} which are pockets of enamel filled with dentine, which are arranged successively like a concertina.V.L. Herridge [https://discovery.ucl.ac.uk/id/eprint/133456/1/133456_Vol.1.pdf Dwarf Elephants on Mediterranean Islands: A Natural Experiment in Parallel Evolution. PhD Thesis, Vol 1.] p. 32 University College London (2010) The number of lophs/lamellae per tooth, as well as the tooth crown height (hypsodonty) is greater in later species.{{Citation |last=Athanassiou |first=Athanassios |title=The Fossil Record of Continental Elephants and Mammoths (Mammalia: Proboscidea: Elephantidae) in Greece |date=2022 |url=https://link.springer.com/10.1007/978-3-030-68398-6_13 |work=Fossil Vertebrates of Greece Vol. 1 |pages=345–391 |editor-last=Vlachos |editor-first=Evangelos |access-date=2023-11-21 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-68398-6_13 |isbn=978-3-030-68397-9|s2cid=245067102 |url-access=subscription }} Elephantids chew using a proal jaw movement involving a forward stroke of the lower jaws, different from the oblique movement using side to side motion of the jaws in more primitive proboscideans.{{Cite journal |last=Saegusa |first=Haruo |date=March 2020 |title=Stegodontidae and Anancus: Keys to understanding dental evolution in Elephantidae |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119302665 |journal=Quaternary Science Reviews |language=en |volume=231 |pages=106176 |doi=10.1016/j.quascirev.2020.106176|bibcode=2020QSRv..23106176S |s2cid=214094348 |url-access=subscription }} The most primitive elephantid Stegotetrabelodon had a long lower jaw with lower tusks and retained permanent premolars similar to many gomphotheres, while modern elephantids lack permanent premolars, with the lower jaw being shortened (brevirostrine) and lower tusks being absent.

{{gallery|File:Tetralophodon -a.jpg|Molar of Tetralophodon, a "tetralophodont gomphothere"|File:Stegotetrabelodon molar.jpg

|Worn molar of Stegotetrabelodon, a primitive elephantid

|File:Loxodonta africana - Molar of an adult.JPG|Molar of a modern African bush elephant (Loxodonta africana)|File:Mammoth fossil tooth - Pleistocene, USA - 23202566565.jpg|Tooth of Mammuthus sp.|width=200|height=180|align=center|title=Comparison of teeth|Mammuthus_sp._sectioned_upper_and_lower_molars_1.jpg|Cross section through elephantid molars}}Elephantids are typically sexually dimorphic, with substantially larger males, with an accelerated growth rate over a longer period of time than females. Elephantidae contains some of the largest known proboscideans, with fully-grown males of some species of mammoths and Palaeoloxodon having average body masses of {{Convert|11|t|lb}} and {{Convert|13|t|lb}} respectively, making them among the largest terrestrial mammals ever. One species of Palaeoloxodon, Palaeoloxodon namadicus, has been suggested to have been possibly the largest land mammal of all time, though this remains speculative due to the fragmentary nature of known remains.{{cite journal |last1=Larramendi |first1=A. |year=2016 |title=Shoulder height, body mass and shape of proboscideans |url=https://www.app.pan.pl/archive/published/app61/app001362014.pdf |journal=Acta Palaeontologica Polonica |volume=61 |doi=10.4202/app.00136.2014 |s2cid=2092950 |doi-access=free}} In comparison to more basal elephantimorphs like gomphotheres, the bodies of elephantids tend to be proportionally shorter from front to back, as well having more elongate limbs with more slender limb bones.{{Cite journal |last1=Bader |first1=Camille |last2=Delapré |first2=Arnaud |last3=Göhlich |first3=Ursula B. |last4=Houssaye |first4=Alexandra |date=November 2024 |title=Diversity of limb long bone morphology among proboscideans: how to be the biggest one in the family |journal=Papers in Palaeontology |language=en |volume=10 |issue=6 |bibcode=2024PPal...10E1597B |doi=10.1002/spp2.1597 |issn=2056-2799 |doi-access=free}}

{{gallery|Jumbo_size_comparison.jpg|Skeleton of an African bush elephant (Loxodonta africana) bull|M. trogontherii skeletal (cropped).png|Skeleton of a steppe mammoth (Mammuthus trogontherii) in front-on (without head) side-on and top-down views|Asian_elephant_size_comparison.jpg|Skeleton of an Asian elephant (Elephas maximus)|Palaeoloxodon_antiquus_size_comparison.png|Skeleton of a straight-tusked elephant (Palaeoloxodon antiquus) bull|Palaeoloxodon falconeri Size Comparison.svg

|Size comparison of Palaeoloxodon falconeri a dwarf elephant species from Sicily, and one of the smallest elephantids known|||width=185|height=|lines=|align=center|title=Gallery of elephantid skeletons}}

{{Further information|Elephant#Behaviour and ecology}}

Living female and juvenile elephants live in matriarchal (female-led) herds of related individuals, with males leaving these groups to live solitarily upon reaching adolescence around 14–15 years of age.{{Cite journal |last1=Neto de Carvalho |first1=Carlos |last2=Belaústegui |first2=Zain |last3=Toscano |first3=Antonio |last4=Muñiz |first4=Fernando |last5=Belo |first5=João |last6=Galán |first6=Jose María |last7=Gómez |first7=Paula |last8=Cáceres |first8=Luis M. |last9=Rodríguez-Vidal |first9=Joaquín |last10=Cunha |first10=Pedro Proença |last11=Cachão |first11=Mario |last12=Ruiz |first12=Francisco |last13=Ramirez-Cruzado |first13=Samuel |last14=Giles-Guzmán |first14=Francisco |last15=Finlayson |first15=Geraldine |date=2021-09-16 |title=First tracks of newborn straight-tusked elephants (Palaeoloxodon antiquus) |journal=Scientific Reports |language=en |volume=11 |issue=1 |page=17311 |bibcode=2021NatSR..1117311N |doi=10.1038/s41598-021-96754-1 |issn=2045-2322 |pmc=8445925 |pmid=34531420}} Evidence has been found that extinct elephantids, including the most primitive elephantid, Stegotetrabelodon, also lived in herds based on footprint tracks.{{Citation |last1=Bibi |first1=Faysal |title=Fossil Trackways of the Baynunah Formation |date=2022 |work=Sands of Time |pages=283–298 |editor-last=Bibi |editor-first=Faysal |url=https://link.springer.com/10.1007/978-3-030-83883-6_17 |access-date=2024-01-18 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-83883-6_17 |isbn=978-3-030-83882-9 |last2=Kraatz |first2=Brian |last3=J. Beech |first3=Mark |last4=Hill |first4=Andrew |editor2-last=Kraatz |editor2-first=Brian |editor3-last=Beech |editor3-first=Mark J. |editor4-last=Hill |editor4-first=Andrew|url-access=subscription }}

{{See also|Elephants}}

File:Palaeoloxodon phylogeny.svg elephantid species, including Palaeoloxodon and mammoths, showing the hybridisation between African forest elephants and Palaeoloxodon, after Palkopoulou et al. 2018]]

File: Comparative view of the human and elephant frame, Benjamin Waterhouse Hawkins, 1860.jpg's A comparative view of the human and animal frame, 1860]]

File:Mammuthus meridionalis.JPG at the French Museum of Natural History]]

Some authors have suggested to classify the family into two subfamilies, Stegotetrabelodontinae, which is monotypic, only containing Stegotetrabelodon, and Elephantinae, containing all other elephantids. Recent genetic research has indicated that Elephas and Mammuthus are more closely related to each other than to Loxodonta, with Palaeoloxodon closely related to Loxodonta. Palaeoloxodon also appears to have received extensive hybridisation with the African forest elephant, and to a lesser extent with mammoths.{{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 |bibcode=2018PNAS..115E2566P |doi=10.1073/pnas.1720554115 |issn=0027-8424 |pmc=5856550 |pmid=29483247 |doi-access=free}}

• Loxodonta (African)
• L. africana African bush elephant
• L. cyclotis African forest elephant
• Elephas (Asiatic)
• E. maximus Asian elephant
• E. m. maximus Sri Lankan elephant
• E. m. indicus Indian elephant
• E. m. sumatranus Sumatran elephant
• E. m. borneensis Borneo elephant

• Elephantidae
• †Stegotetrabelodon (4 species)
• Subfamily Elephantinae
• †Primelephas (2 species)
• Elephas (7+ species)
• †Stegoloxodon (2 species)
• Loxodonta (6 species)
• †Palaeoloxodon (14+ species)
• †Phanagoroloxodon (1 species)
• †Mammuthus (10 species)
• †Stegodibelodon (1 species)
• †Selenetherium (1 species)

Image:ElephEvol.jpg (bottom) to the modern day (top)]]During the Late Miocene, around 10-8 million years ago, the earliest members of the family Elephantidae emerged in Afro-Arabia, having originated from gomphotheres, most likely from members of the genus Tetralophodon.{{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}}{{Citation |last=Sanders |first=William J. |title=Proboscidea from the Baynunah Formation |date=2022 |work=Sands of Time |pages=141–177 |editor-last=Bibi |editor-first=Faysal |url=https://link.springer.com/10.1007/978-3-030-83883-6_10 |access-date=2025-04-17 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-83883-6_10 |isbn=978-3-030-83882-9 |editor2-last=Kraatz |editor2-first=Brian |editor3-last=Beech |editor3-first=Mark J. |editor4-last=Hill |editor4-first=Andrew|url-access=subscription }} The earliest members of the modern genera of Elephantidae appeared during the latest Miocene–early Pliocene around 6-5 million years ago. 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.{{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}} Mammoths then migrated into North America around 1.5 million years ago.{{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}} 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.{{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}} Palaeoloxodon became extinct as part of the Late Pleistocene extinctions, with mammoths only surviving in 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.{{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 |bibcode=2021NatEE...5.1266C |doi=10.1038/s41559-021-01498-w |pmid=34211141 |s2cid=235712060|hdl=10261/249360 |hdl-access=free }}{{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}}

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• Deinotherium
• Embrithopoda
• Eritherium azzouzorum
• Zygolophodon
• Palaeoloxodon namadicus

{{div col end}}

{{Reflist}}

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{{Proboscidea Genera}}

Category:Extant Pliocene first appearances

Category:Mammal families

Category:Proboscideans

Category:Taxa named by John Edward Gray

Category:Elephants