Gomphotherium

Not to be confused with Comphotherium.

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] It is the most diverse genus of gompothere, with over a dozen valid species. The genus is probably paraphyletic.^[3][4]

Gomphotherium Temporal range: Early Miocene–Early Pliocene PreꞒ Ꞓ O S D C P T J K Pg N
Specimen of Gomphotherium productum at the American Museum of Natural History
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 Bunolophodon Vacek, 1877 Tetrabelodon Cope, 1884 Serridentinus Osborn, 1923 Trobelodon Frick, 1933 Ocalientinus Frick, 1933 Tatabelodon Frick, 1933

Description

Skeletal restoration of G. productum (right) and G. steinheimense (left) compared to a human

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 basal elephantimorphs, had an elongated lower jaw which bore tusks.^[6] Species of Gomphotherium are defined by their conservative molar morphology, which includes "trilophed intermediate molars, third molars with three to four loph(id)s, and pretrite half-loph(id)s typically with anterior and posterior accessory conules that form trefoil-patterned enamel loops with wear (simple molar crowns with no accessory conules on the posttrite side of the crown)".^[7]

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] Oxygen and carbon isotopes from G. productum enamel unearthed in the Port of Entry Pit, Oklahoma reveal it fed predominantly on C₃ plants year-round.^[8]

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.^[9] Gomphotherium migrated into Eurasia across the "Gomphotherium land bridge" approximately 19 million years ago.^[10] Gomphotherium underwent rapid evolution after its arrival in Eurasia, reaching its peak diversity during the Early-Middle Miocene.^[10] 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,^[11] and is suggested to be ancestral to later New World gomphothere genera, such as Cuvieronius, Stegomastodon and Rhynchotherium.^[12] Asian populations of Gomphotherium are suggested to have been ancestral to Sinomastodon.^[13] 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.^[13] The last Gomphotherium species disappeared from North America at the beginning of the Pliocene, approximately 5 million years ago.^[11]

Taxonomy

Species

Detail of Gomphotherium skeleton at the American Museum of Natural History

Front view of G. productum. Collected from Clarendon, Texas. At the AMNH.

Over a dozen species of Gomphotherium are considered valid, with over 30 junior synonyms proposed for these taxa.^[14]

• 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
• G. pyrenaicum (Lartet, 1859) Europe, Middle Miocene^[15]

Phylogeny after Wang et al., 2017^[14]

	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"

Cladogram of Elephantiformes after Li et al. 2023, showing a paraphyletic Gomphotheriidae and Gomphotherium.^[16]

Elephantiformes	Phiomia Elephantimorpha Mammutidae (mastodons) Gomphotheres sensu lato Eritreum Gomphotherium annectens Choerolophodontidae Amebelodontidae (shovel tuskers) "Gomphotheriidae" Gomphotherium angustidens Gomphotherium steinheimense Tetralophodon + Elephantidae Gomphotherium sylvaticum Gomphotherium inopinatum Gomphotherium browni Gomphotherium tassyi Gomphotherium productum + American gomphotheres

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