Biology:Gomphotherium

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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 gomphothere, with over a dozen valid species. The genus is probably paraphyletic,[3][4] and ancestral to other gomphothere genera.

Description

Skeletal restoration of G. productum (left) and G. steinheimense (right) 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]

Lower jaws of Gomphotherium angustidens showing elongate mandibular symphysis and lower tusks

Gomphotherium, like most basal elephantimorphs, had an elongated lower jaw (with the elongation specifically being the fused front-most part, the mandibular symphysis) 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] It has been suggested that like other long-jawed elephantimorphs, that the trunk was relatively short in comparison to living elephants, probably not reaching much further than the tips of tusks on the lower jaw.[8]

Ecology

Miniature model of Gomphotherium productum at the Natural History Museum in London

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.[9] Gomphotherium, like other elephantimorphs with long lower jaws, is thought to have used its tusks on the lower jaw to cut through vegetation while it was being held in place by the trunk, which likely had a high level of flexibility similar to living elephants. Its feeding strategy is suggested to have been relatively unspecialised, unlike some other contemporary long-jawed elephantimorphs such as Choerolophodon or the amebelodontid Platybelodon.[10] A 2026 study analyzed the chemical composition of teeth of Gomphotherium angustidens from the Miocene (Langhian) strata from Quinta da Farinheira of Portugal, providing evidence of seasonal changes in its diet and probable evidence of geophagia during fixed times of the year.[11]

Historic 1901 illustration of Gomphotherium angustidens feeding. By Charles R. Knight.

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.[12] Gomphotherium migrated into Eurasia across the "Gomphotherium land bridge" approximately 19 million years ago.[13] Gomphotherium underwent rapid evolution after its arrival in Eurasia, reaching its peak diversity during the Early-Middle Miocene.[13] Gomphotherium has been posited to be paraphyletic and the ancestor of later gomphothere genera, including the "tetralophodont gomphotheres", such as Tetralophodon (which is suggested to have a close relationship with the species G. steinheimense) which are probably ancestral to elephantids.[3] Gomphotherium annectens has been suggested to be the ancestor of the stegodontids.[14] Gomphotherium first arrived in North America during the mid-Miocene, approximately 16-15 million years ago,[15] and is suggested to be ancestral to later New World gomphothere genera, such as Cuvieronius, Stegomastodon and Rhynchotherium.[16] Asian populations of Gomphotherium are suggested to have been ancestral to Sinomastodon.[17] Gomphotherium became extinct in northern China during the Middle Miocene around 13 million years ago.[10] The youngest remains attributed to Gomphotherium in Africa date to around 13-10 million years ago.[7] 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.[17] The last Gomphotherium species disappeared from North America at the beginning of the Pliocene, approximately 5 million years ago.[15]

Taxonomy

Species

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.[18]

  • 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[19]

Phylogeny after Wang et al., 2017[18]

 
 

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

 
 
 
 
 
 
 
 
 
 
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Cladogram of Elephantiformes after Li et al. 2023, showing a paraphyletic Gomphotheriidae and Gomphotherium.[20]

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

 
 
 
 
 
 
 
 
 
 
 

References

  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. doi:10.1016/j.quaint.2014.06.036. ISSN 1040-6182. Bibcode: 2014QuInt.354..122W. https://www.researchgate.net/publication/263930469. 
  2. ↑ Palmer, T. S.; Merriam, C. H. (1904). Index generum mammalium: a list of the genera and families of mammals. Government Printing Office, Washington.
  3. ↑ 3.0 3.1 3.2 Wu, Yan; Deng, Tao; Hu, Yaowu; Ma, Jiao; Zhou, Xinying; Mao, Limi; Zhang, Hanwen; Ye, Jie et al. (16 May 2018). "A grazing Gomphotherium in Middle Miocene Central Asia, 10 million years prior to the origin of the Elephantidae" (in en). Scientific Reports 8 (1): 7640. doi:10.1038/s41598-018-25909-4. ISSN 2045-2322. PMID 29769581. Bibcode: 2018NatSR...8.7640W. 
  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" (in en). iScience 25 (1). doi:10.1016/j.isci.2021.103559. PMID 34988402. Bibcode: 2022iSci...25j3559B. 
  5. ↑ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans". Acta Palaeontologica Polonica 61. doi:10.4202/app.00136.2014. https://www.app.pan.pl/archive/published/app61/app001362014.pdf. 
  6. ↑ Mothé, Dimila; Ferretti, Marco P.; Avilla, Leonardo S. (2016-01-12). Beatty, Brian Lee. ed. "The Dance of Tusks: Rediscovery of Lower Incisors in the Pan-American Proboscidean Cuvieronius hyodon Revises Incisor Evolution in Elephantimorpha" (in en). PLOS ONE 11 (1). doi:10.1371/journal.pone.0147009. ISSN 1932-6203. PMID 26756209. Bibcode: 2016PLoSO..1147009M. 
  7. ↑ 7.0 7.1 Sanders, William J. (2023-07-07) (in en). Evolution and Fossil Record of African Proboscidea (1 ed.). Boca Raton: CRC Press. pp. 94, 106-107. doi:10.1201/b20016. ISBN 978-1-315-11891-8. https://www.taylorfrancis.com/books/9781315118918. 
  8. ↑ Nabavizadeh, Ali (2024-10-08). "Of tusks and trunks: A review of craniofacial evolutionary anatomy in elephants and extinct Proboscidea" (in en). The Anatomical Record. doi:10.1002/ar.25578. ISSN 1932-8486. PMID 39380178. https://anatomypubs.onlinelibrary.wiley.com/doi/10.1002/ar.25578. 
  9. ↑ Fox, David L.; Fisher, Daniel C. (1 June 2001). "Stable Isotope Ecology of a Late Miocene Population of Gomphotherium productus (Mammalia, Proboscidea) from Port of Entry Pit, Oklahoma, USA" (in en). PALAIOS 16 (3): 279–293. doi:10.1669/0883-1351(2001)016<0279:SIEOAL>2.0.CO;2. ISSN 0883-1351. Bibcode: 2001Palai..16..279F. https://bioone.org/journals/palaios/volume-16/issue-3/0883-1351_2001_016_0279_SIEOAL_2.0.CO_2/Stable-Isotope-Ecology-of-a-Late-Miocene-Population-of-Gomphotherium/10.1669/0883-1351(2001)016%3C0279:SIEOAL%3E2.0.CO;2.short. Retrieved 2 July 2024. 
  10. ↑ 10.0 10.1 Li, Chunxiao; Deng, Tao; Wang, Yang; Sun, Fajun; Wolff, Burt; Jiangzuo, Qigao; Ma, Jiao; Xing, Luda et al. (2023-11-28), "The trunk replaces the longer mandible as the main feeding organ in elephant evolution", eLife 12, doi:10.7554/eLife.90908.1, https://elifesciences.org/articles/90908, retrieved 2024-05-29 
  11. ↑ Coimbra, Rute; de Winter, Niels; Ríos, Maria; Bernardino, Rui; Estraviz-López, Darío; Lohmann, Priscila; Martino, Roberta; Grandal-d'Anglade, Aurora et al. (2026-03-10). "Detection of dietary stress and geophagic behaviour forced by dry seasons in Miocene Gomphotherium" (in en). Biogeosciences 23 (5): 1833–1858. doi:10.5194/bg-23-1833-2026. ISSN 1726-4189. https://bg.copernicus.org/articles/23/1833/2026/. 
  12. ↑ Wang, Shi-Qi; Li, Yu; Duangkrayom, Jaroon; Yang, Xiang-Wen; He, Wen; Chen, Shan-Qin (2017-05-04). "A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia" (in en). Journal of Vertebrate Paleontology 37 (3). doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. Bibcode: 2017JVPal..37E8284W. https://www.tandfonline.com/doi/full/10.1080/02724634.2017.1318284. 
  13. ↑ 13.0 13.1 Li, Chunxiao; Wang, Shi-Qi; Yang, Qing (2022-05-26). "Discovery of a primitive Gomphotherium from the Early Miocene of northern China and its biochronology and palaeobiogeography significance" (in en). Historical Biology: 1–9. doi:10.1080/08912963.2022.2077106. ISSN 0891-2963. https://www.tandfonline.com/doi/full/10.1080/08912963.2022.2077106. 
  14. ↑ Wang, Shi-Qi; Li, Chunxiao; Li, Yan; Zhang, Xiaoxiao (2023-03-01). "Gomphotheres from Linxia Basin, China, and their significance in biostratigraphy, biochronology, and paleozoogeography" (in en). Palaeogeography, Palaeoclimatology, Palaeoecology 613. doi:10.1016/j.palaeo.2023.111405. https://linkinghub.elsevier.com/retrieve/pii/S0031018223000238. 
  15. ↑ 15.0 15.1 MacFadden, Bruce J.; Morgan, Gary S.; Jones, Douglas S.; Rincon, Aldo F. (March 2015). "Gomphothere proboscidean ( Gomphotherium ) from the late Neogene of Panama" (in en). Journal of Paleontology 89 (2): 360–365. doi:10.1017/jpa.2014.31. ISSN 0022-3360. Bibcode: 2015JPal...89..360M. 
  16. ↑ Spencer LG 2022. The last North American gomphotheres. N Mex Mus Nat Hist Sci. 88:45–58.
  17. ↑ 17.0 17.1 Wang, Shi-Qi; Ji, Xue-Ping; Jablonski, Nina G.; Su, Denise F.; Ge, Jun-Yi; Ding, Chang-Fen; Yu, Teng-Song; Li, Wen-Qi et al. (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" (in en). Journal of Mammalian Evolution 23 (2): 155–173. doi:10.1007/s10914-015-9311-z. ISSN 1064-7554. http://link.springer.com/10.1007/s10914-015-9311-z. 
  18. ↑ 18.0 18.1 Wang, Shi-Qi; Li, Yu; Duangkrayom, Jaroon; Yang, Xiang-Wen; He, Wen; Chen, Shan-Qin (2017-05-04). "A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia". Journal of Vertebrate Paleontology 37 (3). doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. Bibcode: 2017JVPal..37E8284W. 
  19. ↑ Göhlich, Ursula B. (2010). "The Proboscidea (Mammalia) from the Miocene of Sandelzhausen (southern Germany)". Paläontologische Zeitschrift 84 (1): 163–204. doi:10.1007/s12542-010-0053-1. Bibcode: 2010PalZ...84..163G. 
  20. ↑ Li, Chunxiao; Deng, Tao; Wang, Yang; Sun, Fajun; Wolff, Burt; Jiangzuo, Qigao; Ma, Jiao; Xing, Luda et al. (2023-11-28), "Longer mandible or nose? Co-evolution of feeding organs in early elephantiforms", eLife 12, doi:10.7554/eLife.90908.1, https://elifesciences.org/reviewed-preprints/90908v1, retrieved 2024-05-29 


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