Gomphotherium Annectens
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''Gomphotherium'' (; "nail beast" for its double set of straight tusks) is an extinct genus of
gomphothere Gomphotheres are an extinct group of proboscideans related to modern elephants. First appearing in Africa during the Oligocene, they dispersed into Eurasia and North America during the Miocene and arrived in South America during the Pleistocene a ...
proboscidea Proboscidea (; , ) is a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Three l ...
n from the
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
of Eurasia, Africa and North America. It is the most diverse genus of gompothere, with over a dozen valid species. The genus is probably
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
.


Description

Most species of ''Gomphotherium'' were similar in size to the
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living ''Elephas'' species. It is the largest living land animal in Asia and the second largest living Elephantidae, elephantid in the world. It is char ...
, with ''G. productum'' (known from a 35-year-old male) measuring tall and weighing . The largest species ''G. steinheimense'', known from a complete 37-year-old male found in
Mühldorf Mühldorf am Inn (, ; Central Bavarian: ''Muihdorf am Inn'') is a Town#Germany, town in Bavaria, Germany, and the capital of the Mühldorf (district), district Mühldorf on the river Inn (river), Inn. It is located at , and had a population of ab ...
, Germany, measured up to tall and weighed . ''Gomphotherium'', like most basal elephantimorphs, had an elongated lower jaw which bore tusks. 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)".


Ecology

Most species of ''Gomphotherium'' are inferred to have been
browsers Browse, browser, or browsing may refer to: Computing *Browser service, a feature of Microsoft Windows to browse shared network resources *Code browser, a program for navigating source code *File browser or file manager, a program used to manage f ...
or mixed feeders, but specimens of ''G''. ''steinheimense'' from China are suggested to have been grazers. 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.


Evolution

''Gomphotherium'' likely originated in Africa during the late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
-early
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
. The oldest remains of ''Gomphotherium'' are known from Africa, dating to approximately 19.5 million years ago''. Gomphotherium'' migrated into Eurasia across the " ''Gomphotherium'' land bridge" approximately 19 million years ago. ''Gomphotherium'' underwent rapid evolution after its arrival in Eurasia, reaching its peak diversity during the Early-Middle Miocene. ''Gomphotherium'' has been posited to be
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
and the ancestor of later gomphothere genera, including the "tetralophodont gomphotheres" such as ''
Tetralophodon ''Tetralophodon'' ("four-ridged tooth") is an extinct genus of "tetralophodont gomphothere" belonging to the superfamily Elephantoidea, known from the Miocene of Afro-Eurasia. Taxonomy and evolution The genus ''Tetralophodon'' (meaning "four-rid ...
'' which are probably ancestral to stegodontids and elephantids. ''Gomphotherium'' first arrived in North America during the mid-Miocene, approximately 16-15 million years ago, and is suggested to be ancestral to later New World gomphothere genera, such as ''
Cuvieronius ''Cuvieronius'' is an extinct New World genus of gomphothere which ranged from southern North America to western South America during the Pleistocene epoch. Reaching a shoulder height of and a body mass of , it was on average shorter but compara ...
'', ''
Stegomastodon ''Stegomastodon'' ('roof breast tooth') is an extinct genus of gomphotheres. It ranged throughout North America from the Pliocene (early Blancan ~4 Ma), to the Early Pleistocene (early Irvingtonian, ~1.2 Ma). The former South American species h ...
'' and ''
Rhynchotherium ''Rhynchotherium'' is an extinct genus of proboscidea endemic to North America and Central America during the Miocene through Pliocene from 13.650 to 3.6 Annum, Ma, living for approximately . This gomphothere had two tusks and may have evolved f ...
.'' Asian populations of ''Gomphotherium'' are suggested to have been ancestral to ''
Sinomastodon ''Sinomastodon'' ("Chinese mastodont") is an extinct gomphothere genus (of order Proboscidea) known from the Late Miocene to Early Pleistocene of Asia, including China, Japan, Thailand, Myanmar, Indonesia and probably Kashmir. Extinct animals o ...
''. 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. The last ''Gomphotherium'' species disappeared from North America at the beginning of the Pliocene, approximately 5 million years ago.


Taxonomy


Species

Over a dozen species of ''Gomphotherium'' are considered valid, with over 30 junior synonyms proposed for these taxa. * ''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 Phylogeny after Wang ''et al.'', 2017 Cladogram of Elephantiformes after Li et al. 2023, showing a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
Gomphotheriidae Gomphotheres are an extinct group of proboscideans related to modern elephants. First appearing in Africa during the Oligocene, they dispersed into Eurasia and North America during the Miocene and arrived in South America during the Pleistocene ...
and ''Gomphotherium''. File:Gomphotherium NT small.jpg, ''Gomphotherium angustidens'' File:Gomphotherium productum 3.JPG, ''Gomphotherium productum'' File:Knight Gomphotherium.jpg, ''G. angustidens'' by Charles R. Knight File:Gomphotherium angustidens 2.jpg, ''G. angustidens'' skeleton File:Gomphotherium angustidens mandibles.jpg,
Ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the ovum, egg to adult. The term can also be used to refer to t ...


References

{{Taxonbar, from=Q132604 Miocene mammals of Africa Miocene mammals of Europe Gomphotheres Miocene proboscideans Pliocene proboscideans Zanclean extinctions Miocene mammals of North America White River Fauna Prehistoric placental genera Fossil taxa described in 1837 Miocene mammals of Asia Pliocene mammals of North America