Gomphotherium

Gomphotherium; Temporal range: Early Miocene–Early Pliocene PreꞒ Ꞓ O S D C P T J K Pg N
	Specimen of G. productum (AMNH)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Proboscidea
Family:	†Gomphotheriidae
Genus:	†Gomphotherium; Burmeister, 1837
Type species
	Gomphotherium angustidens; (Cuvier, 1817)
Species
	See text
Synonyms
	Trilophodon; Falconer and Cautley, 1846; Tetrabelodon; Cope, 1884; Serridentinus; Osborn, 1923; Tatabelodon; Frick 1933;

Gomphotherium (/ˌɡɒmfəˈθɪəriəm/; "nail beast" for its double set of straight tusks) is an extinct genus of gomphothere proboscidean from the Neogene of Eurasia, Africa and North America.^[1]^[2] The genus is probably paraphyletic.^[3]^[4]

Description

Most species of Gomphotherium were similar in size to the Asian elephant, with G. productum (known from a 35-year-old male) measuring 2.51 m (8 ft 3 in) tall and weighing 4.6 t (4.5 long tons; 5.1 short tons). The largest species G. steinheimense, known from a complete 37-year-old male found in Mühldorf, Germany, measured up to 3.17 m (10.4 ft) tall and weighed 6.7 t (6.6 long tons; 7.4 short tons).^[5]

Gomphotherium, like most primitive elephantimorphs, had an elongated lower jaw which bore tusks.^[6]

Ecology

Most species of Gomphotherium are inferred to have been browsers or mixed feeders, but specimens of G. steinheimense from China are suggested to have been grazers.^[3]

Evolution

Gomphotherium likely originated in Africa during the late Oligocene-early Miocene. The oldest remains of Gomphotherium are known from Africa, dating to approximately 19.5 million years ago.^[7] Gomphotherium migrated into Eurasia across the "Gomphotherium land bridge" approximately 19 million years ago.^[8] Gomphotherium underwent rapid evolution after its arrival in Eurasia, reaching its peak diversity during the Early-Middle Miocene.^[8] Gomphotherium has been posited to be paraphyletic and the ancestor of later gomphothere genera, including the "tetralophodont gomphotheres" such as Tetralophodon which are probably ancestral to stegodontids and elephantids.^[3] Gomphotherium first arrived in North America during the mid-Miocene, approximately 16-15 million years ago,^[9] and is suggested to be ancestral to later New World gomphothere genera, such as Cuvieronius, Stegomastodon and Rhynchotherium.^[10] Asian populations of Gomphotherium are suggested to have been ancestral to Sinomastodon.^[11] The last European species of Gomphotherium became extinct at the beginning of the Late Miocene, around the start of MN9, approximately 10 million years ago.^[11] The last Gomphotherium species disappeared from North America at the beginning of the Pliocene, approximately 5 million years ago.^[9]

Taxonomy

The following cladogram shows the placement of the genus Gomphotherium among other proboscideans, based on hyoid characteristics:^[12]

Mammut americanum (American mastodon)

Gomphotherium sp.

Stegodon zdanskyi

Loxodonta africana (African elephant)

	Elephas maximus (Asian elephant)

	Mammuthus columbi (Columbian mammoth)

Species

Over a dozen species of Gomphotherium are considered valid, with over 30 junior synonyms proposed for these taxa.^[13]

G. hannibali Welcomme, 1994 Europe, Early Miocene
G. annectens (Matsumoto, 1925) Japan, Early Miocene
G. cooperi (Osborn, 1932) Asia, Early Miocene
G. sylvaticum Tassy, 1985 Europe, Early Miocene
G. libycum (Fourtau, 1918) Egypt, Early Miocene
G. inopinatum (Borissiak and Belyaeva, 1928) China, late Early Miocene-Early middle Miocene
G. mongoliense (Osborn, 1924) Mongolia, late Early Miocene-Early middle Miocene
G. angustidens (Cuvier, 1817) (type) Europe, Middle Miocene
G. subtapiroideum (Schlesinger, 1917) Europe, Early-Middle Miocene
G. tassyi Wang, Li, Duangkrayom, Yang, He & Chen, 2017 China, Middle Miocene
G. browni (Osborn, 1926) Pakistan, Middle Miocene
G. steinheimense (Klahn, 1922) Europe, China, Middle-Late Miocene
G. productum (Cope, 1874) North America Middle Miocene-Early Pliocene

Phylogeny after Wang et al. 2017^[13]

Phiomia serridens

Eritreum melakeghebrekristosi

Gomphotherium sp. (Mwiti)

Gomphotherium hannibali

Gomphotherium annectens

Gomphotherium cooperi

Gomphotherium sylvaticum

	Gomphotherium libycum

	Gomphotherium pygmaeus

Gomphotherium inopinatum

Gomphotherium mongoliense

	Gomphotherium angustidens (s. s.)

	Gomphotherium connexum

	Gomphotherium subtapiroideum

	Gomphotherium tassyi

Gomphotherium wimani

	Gomphotherium browni

	Gomphotherium productum

	Gomphotherium steinheimense

Outgroups

"G. annectens group"

"G. angustidens group"

"Derived Gomphotherium"

Gomphotherium angustidens
Gomphotherium productum
G. angustidens by Charles R. Knight
G. angustidens skeleton

References

^ Wang, Wei; Liao, Wei; Li, Dawei; Tian, Feng (1 July 2014). "Early Pleistocene large-mammal fauna associated with Gigantopithecus at Mohui Cave, Bubing Basin, South China". Quaternary International. 354: 122–130. Bibcode:2014QuInt.354..122W. doi:10.1016/j.quaint.2014.06.036. ISSN 1040-6182.
^ Palmer, T. S.; Merriam, C. H. (1904). Index generum mammalium: a list of the genera and families of mammals. Government Printing Office, Washington.
^ ^a ^b ^c Wu, Yan; Deng, Tao; Hu, Yaowu; Ma, Jiao; Zhou, Xinying; Mao, Limi; Zhang, Hanwen; Ye, Jie; Wang, Shi-Qi (16 May 2018). "A grazing Gomphotherium in Middle Miocene Central Asia, 10 million years prior to the origin of the Elephantidae". Scientific Reports. 8 (1): 7640. Bibcode:2018NatSR...8.7640W. doi:10.1038/s41598-018-25909-4. ISSN 2045-2322. PMC 5956065. PMID 29769581.
^ Baleka, Sina; Varela, Luciano; Tambusso, P. Sebastián; Paijmans, Johanna L.A.; Mothé, Dimila; Stafford, Thomas W.; Fariña, Richard A.; Hofreiter, Michael (January 2022). "Revisiting proboscidean phylogeny and evolution through total evidence and palaeogenetic analyses including Notiomastodon ancient DNA". iScience. 25 (1): 103559. Bibcode:2022iSci...25j3559B. doi:10.1016/j.isci.2021.103559. PMC 8693454. PMID 34988402.
^ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi:10.4202/app.00136.2014. S2CID 2092950.
^ Mothé, Dimila; Ferretti, Marco P.; Avilla, Leonardo S. (12 January 2016). Beatty, Brian Lee (ed.). "The Dance of Tusks: Rediscovery of Lower Incisors in the Pan-American Proboscidean Cuvieronius hyodon Revises Incisor Evolution in Elephantimorpha". PLOS ONE. 11 (1): e0147009. Bibcode:2016PLoSO..1147009M. doi:10.1371/journal.pone.0147009. ISSN 1932-6203. PMC 4710528. PMID 26756209.
^ Wang, Shi-Qi; Li, Yu; Duangkrayom, Jaroon; Yang, Xiang-Wen; He, Wen; Chen, Shan-Qin (4 May 2017). "A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia". Journal of Vertebrate Paleontology. 37 (3): e1318284. Bibcode:2017JVPal..37E8284W. doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. S2CID 90593535.
^ ^a ^b Li, Chunxiao; Wang, Shi-Qi; Yang, Qing (26 May 2022). "Discovery of a primitive Gomphotherium from the Early Miocene of northern China and its biochronology and palaeobiogeography significance". Historical Biology: 1–9. doi:10.1080/08912963.2022.2077106. ISSN 0891-2963.
^ ^a ^b MacFadden, Bruce J.; Morgan, Gary S.; Jones, Douglas S.; Rincon, Aldo F. (March 2015). "Gomphothere proboscidean ( Gomphotherium ) from the late Neogene of Panama". Journal of Paleontology. 89 (2): 360–365. Bibcode:2015JPal...89..360M. doi:10.1017/jpa.2014.31. ISSN 0022-3360. S2CID 52093950.
^ Spencer LG 2022. The last North American gomphotheres. N Mex Mus Nat Hist Sci. 88:45–58.
^ ^a ^b Wang, Shi-Qi; Ji, Xue-Ping; Jablonski, Nina G.; Su, Denise F.; Ge, Jun-Yi; Ding, Chang-Fen; Yu, Teng-Song; Li, Wen-Qi; Duangkrayom, Jaroon (June 2016). "The Oldest Cranium of Sinomastodon (Proboscidea, Gomphotheriidae), Discovered in the Uppermost Miocene of Southwestern China: Implications for the Origin and Migration of This Taxon". Journal of Mammalian Evolution. 23 (2): 155–173. doi:10.1007/s10914-015-9311-z. ISSN 1064-7554.
^ Shoshani, J.; Tassy, P. (2005). "Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior". Quaternary International. 126–128: 5–20. Bibcode:2005QuInt.126....5S. doi:10.1016/j.quaint.2004.04.011.
^ ^a ^b Wang, Shi-Qi; Li, Yu; Duangkrayom, Jaroon; Yang, Xiang-Wen; He, Wen; Chen, Shan-Qin (4 May 2017). "A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia". Journal of Vertebrate Paleontology. 37 (3): e1318284. Bibcode:2017JVPal..37E8284W. doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. S2CID 90593535.

[1] Wang, Wei; Liao, Wei; Li, Dawei; Tian, Feng (1 July 2014). "Early Pleistocene large-mammal fauna associated with Gigantopithecus at Mohui Cave, Bubing Basin, South China". Quaternary International. 354: 122–130. Bibcode:2014QuInt.354..122W. doi:10.1016/j.quaint.2014.06.036. ISSN 1040-6182.

[2] Palmer, T. S.; Merriam, C. H. (1904). Index generum mammalium: a list of the genera and families of mammals. Government Printing Office, Washington.

[:0-3] Wu, Yan; Deng, Tao; Hu, Yaowu; Ma, Jiao; Zhou, Xinying; Mao, Limi; Zhang, Hanwen; Ye, Jie; Wang, Shi-Qi (16 May 2018). "A grazing Gomphotherium in Middle Miocene Central Asia, 10 million years prior to the origin of the Elephantidae". Scientific Reports. 8 (1): 7640. Bibcode:2018NatSR...8.7640W. doi:10.1038/s41598-018-25909-4. ISSN 2045-2322. PMC 5956065. PMID 29769581.

[4] Baleka, Sina; Varela, Luciano; Tambusso, P. Sebastián; Paijmans, Johanna L.A.; Mothé, Dimila; Stafford, Thomas W.; Fariña, Richard A.; Hofreiter, Michael (January 2022). "Revisiting proboscidean phylogeny and evolution through total evidence and palaeogenetic analyses including Notiomastodon ancient DNA". iScience. 25 (1): 103559. Bibcode:2022iSci...25j3559B. doi:10.1016/j.isci.2021.103559. PMC 8693454. PMID 34988402.

[proboscideans-5] Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi:10.4202/app.00136.2014. S2CID 2092950.

[6] Mothé, Dimila; Ferretti, Marco P.; Avilla, Leonardo S. (12 January 2016). Beatty, Brian Lee (ed.). "The Dance of Tusks: Rediscovery of Lower Incisors in the Pan-American Proboscidean Cuvieronius hyodon Revises Incisor Evolution in Elephantimorpha". PLOS ONE. 11 (1): e0147009. Bibcode:2016PLoSO..1147009M. doi:10.1371/journal.pone.0147009. ISSN 1932-6203. PMC 4710528. PMID 26756209.

[7] Wang, Shi-Qi; Li, Yu; Duangkrayom, Jaroon; Yang, Xiang-Wen; He, Wen; Chen, Shan-Qin (4 May 2017). "A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia". Journal of Vertebrate Paleontology. 37 (3): e1318284. Bibcode:2017JVPal..37E8284W. doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. S2CID 90593535.

[:3-8] Li, Chunxiao; Wang, Shi-Qi; Yang, Qing (26 May 2022). "Discovery of a primitive Gomphotherium from the Early Miocene of northern China and its biochronology and palaeobiogeography significance". Historical Biology: 1–9. doi:10.1080/08912963.2022.2077106. ISSN 0891-2963.

[:1-9] MacFadden, Bruce J.; Morgan, Gary S.; Jones, Douglas S.; Rincon, Aldo F. (March 2015). "Gomphothere proboscidean ( Gomphotherium ) from the late Neogene of Panama". Journal of Paleontology. 89 (2): 360–365. Bibcode:2015JPal...89..360M. doi:10.1017/jpa.2014.31. ISSN 0022-3360. S2CID 52093950.

[10] Spencer LG 2022. The last North American gomphotheres. N Mex Mus Nat Hist Sci. 88:45–58.

[:4-11] Wang, Shi-Qi; Ji, Xue-Ping; Jablonski, Nina G.; Su, Denise F.; Ge, Jun-Yi; Ding, Chang-Fen; Yu, Teng-Song; Li, Wen-Qi; Duangkrayom, Jaroon (June 2016). "The Oldest Cranium of Sinomastodon (Proboscidea, Gomphotheriidae), Discovered in the Uppermost Miocene of Southwestern China: Implications for the Origin and Migration of This Taxon". Journal of Mammalian Evolution. 23 (2): 155–173. doi:10.1007/s10914-015-9311-z. ISSN 1064-7554.

[12] Shoshani, J.; Tassy, P. (2005). "Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior". Quaternary International. 126–128: 5–20. Bibcode:2005QuInt.126....5S. doi:10.1016/j.quaint.2004.04.011.

[:2-13] Wang, Shi-Qi; Li, Yu; Duangkrayom, Jaroon; Yang, Xiang-Wen; He, Wen; Chen, Shan-Qin (4 May 2017). "A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia". Journal of Vertebrate Paleontology. 37 (3): e1318284. Bibcode:2017JVPal..37E8284W. doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. S2CID 90593535.

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