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===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}}</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>
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