Biology:Gomphothere

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Short description: Extinct family of proboscidean mammals

Gomphothere
Temporal range: 28–0.01 Ma
Late Oligocene - Holocene
Gomphotherium productum.jpg
Specimen of Gomphotherium productum at the AMNH
Stegomastodon CCB.JPG
Notiomastodon platensis Centro Cultural del Bicentenario de Santiago del Estero in Argentina
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Superfamily: †Gomphotherioidea
Family: †Gomphotheriidae
(Hay, 1922) A. Cabrera 1929
Genera

Gomphotheres are an extinct group of proboscideans related to modern elephants. They were widespread across Afro-Eurasia and North America during the Miocene and Pliocene epochs and dispersed into South America during the Pleistocene as part of the Great American Interchange. Gomphotheres are a paraphyletic group that is ancestral to Elephantidae, which contains modern elephants, as well as Stegodontidae. While most famous forms such as Gomphotherium had long lower jaws with tusks, which is the ancestral condition for the group, some later members developed shortened (brevirostrine) lower jaws with either vestigial or no lower tusks, looking very similar to modern elephants, an example of parallel evolution, which outlasted the long-jawed gomphotheres. By the end of the Early Pleistocene, gomphotheres became extinct in Afro-Eurasia, with the last two genera, Cuvieronius ranging from southern North America to western South America, and Notiomastodon having a wide range over most of South America until the end of the Pleistocene around 12,000 years ago, when they became extinct following the arrival of humans.

The name "gomphothere" comes from Ancient Greek γόμφος (gómphos), "peg, pin; wedge; joint" plus θηρίον (theríon), "beast".

Description

Life restoration of Gomphotherium
Life restoration of Cuvieronius, a brevirostrine gomphothere

Gomphotheres differed from elephants in their tooth structure, particularly the chewing surfaces on the molar teeth. The teeth are considered to be bunodont, that is, having rounded cusps.[1] They are thought to have chewed differently from modern elephants, using an oblique movement (combining back to front and side to side motion) over the teeth rather than the proal movement (a forwards stroke from the back to the front of the lower jaws) used by modern elephants and stegodontids.[2] Earlier gomphotheres had lower jaws with an elongate mandibular symphysis and lower tusks, the primitive condition for members of Elephantimorpha. Later members developed shortened (brevirostrine) lower jaws and/or vestigial or no lower tusks, a convergent process that occurred multiple times among gomphotheres, as well as other members of Elephantimorpha.[3] The incisors and long lower jaws of primitive gomphotheres were likely used for cutting vegetation, while brevirostrine gomphotheres relied on their trunks to acquire food similar to modern elephants.[4]

Comparison of teeth

  • Largely unworn molar of Gomphotherium angustidens, a "trilophodont gomphothere"

  • Worn molar of Gomphotherium angustidens

  • Molar of Tetralophodon, a "tetralophodont gomphothere "

  • Molar of a modern African elephant (Loxodonta) for comparison

Taxonomy

"Gomphotheres" are assigned to their own family, Gomphotheriidae, but are widely agreed to be a paraphyletic group. The families Choerolophodontidae and Amebelodontidae (which includes "shovel tuskers" with flattened lower tusks like Platybelodon) are sometimes considered gomphotheres sensu lato,[5][6][7] though some authors argue that Amebelodontidae should be sunk into Gomphotheriidae.[8] Gomphotheres are divided into two informal groups, "trilophodont gomphotheres", and "tetralophodont gomphotheres". "Tetralophodont gomphotheres" are distinguished from "trilophodont gomphotheres" by the presence of four ridges on the fourth premolar and on the first and second molars, rather than the three present in trilophodont gomphotheres.[5] Some authors choose to exclude "tetralophodont gomphotheres" from Gomphotheriidae, and instead assign them to the group Elephantoidea.[5] "Tetralophodont gomphotheres" are thought to have evolved from "trilophodont gomphotheres", and are suggested to be ancestral to Elephantidae, the group which contains modern elephants, as well as Stegodontidae.[9]

Ecology

Gomphotheres are generally supposed to have been flexible feeders, with the various species having differing browsing, mixed feeding and grazing diets, with the dietary preference of individual species and populations being shaped by local factors such as climactic conditions and competition.[10] Analysis of the tusks of a male Notiomastodon individual suggest that it underwent musth, similar to modern elephants.[11] Notiomastiodon is also suggested to have lived in social family groups, like modern elephants.[12]

Evolutionary history

Gomphotheres originated in Afro-Arabia during the mid-Oligocene, with remains from the Shumaysi Formation in Saudi Arabia dating to around 29-28 million years ago. Gomphotheres were uncommon in Afro-Arabia during the Oligocene.[13] Gomphotheres arrived in Eurasia after the connection of Afro-Arabia and Eurasia during the Early Miocene around 19 million years ago,[14] in what is termed the "Proboscidean Datum Event". Gomphotherium arrived in North America around 16 million years ago,[15] and is suggested to be the ancestor of later New World gomphothere genera.[16] "Trilophodont gomphotheres" dramatically declined during the Late Miocene, likely due to the increasing C4 grass-dominated habitats,[14] while during the Late Miocene "tetralophodont gomphotheres" were abundant and widespread in Eurasia, where they represented the dominant group of proboscideans.[17] All trilophodont gomphotheres, with the exception of the Asian Sinomastodon, became extinct in Eurasia by the beginning of the Pliocene,[18] along with the global extinction of the "shovel tusker" amebelodontids.[19] The last gomphotheres in Africa, represented by the "tetralophodont gomphothere" genus Anancus, became extinct around the end of the Pliocene and beginning of the Pleistocene.[20] The New World gomphothere genera Notiomastodon and Cuvieronius dispersed into South America during the Pleistocene, around or after 2.5 million years ago as part of the Great American Biotic Interchange due to the formation of the Isthmus of Panama.[21] The last gomphothere native to Europe, Anancus arvernensis[22] became extinct during the Early Pleistocene, around 1.6–2 million years ago[23][24] Sinomastodon became extinct at the end of the Early Pleistocene, around 800,000 years ago.[25]

The extinction of gomphotheres in Afro-Eurasia has generally been supposed to be the result the expansion of Elephantidae and Stegodon.[18][26] The morphology of elephantid molars being more efficient than gomphotheres in consuming grass, which became more abundant during the Pliocene and Pleistocene epochs.[26] In the New World, gomphotheres did not become extinct until shortly after the arrival of humans to the Americas, approximately 12,000 years ago. Bones of the last gomphothere genera, Cuvieronius and Notiomastodon, dating to shortly before their extinction have been found associated with human artifacts, suggesting that hunting may have played a role in their extinction.[21]

References

  1. ↑ Buckley, Michael; Recabarren, Omar P.; Lawless, Craig; García, Nuria; Pino, Mario (November 2019). "A molecular phylogeny of the extinct South American gomphothere through collagen sequence analysis" (in en). Quaternary Science Reviews 224: 105882. doi:10.1016/j.quascirev.2019.105882. Bibcode: 2019QSRv..22405882B. 
  2. ↑ Saegusa, Haruo (March 2020). "Stegodontidae and Anancus: Keys to understanding dental evolution in Elephantidae" (in en). Quaternary Science Reviews 231: 106176. doi:10.1016/j.quascirev.2020.106176. Bibcode: 2020QSRv..23106176S. https://linkinghub.elsevier.com/retrieve/pii/S0277379119302665. 
  3. ↑ Mothé, Dimila; Ferretti, Marco P.; Avilla, Leonardo S. (12 January 2016). "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. doi:10.1371/journal.pone.0147009. PMID 26756209. Bibcode: 2016PLoSO..1147009M. 
  4. ↑ Li, Chunxiao; Deng, Tao; Wang, Yang; Sun, Fajun; Wolff, Burt; Jiangzuo, Qigao; Ma, Jiao; Xing, Luda et al. (2023-08-16) (in en). Longer mandible or nose? Co-evolution of feeding organs in early elephantiforms (Report). Paleontology. doi:10.1101/2023.08.15.553347. http://biorxiv.org/lookup/doi/10.1101/2023.08.15.553347. 
  5. ↑ 5.0 5.1 5.2 Shoshani, J.; Tassy, P. (2005). "Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior". Quaternary International 126-128: 5–20. doi:10.1016/j.quaint.2004.04.011. Bibcode: 2005QuInt.126....5S. 
  6. ↑ Wang, Shi-Qi; Deng, Tao; Ye, Jie; He, Wen; Chen, Shan-Qin (2016). "Morphological and ecological diversity of Amebelodontidae (Proboscidea, Mammalia) revealed by a Miocene fossil accumulation of an upper-tuskless proboscidean". Systematic Palaeontology 15 (8): 601–615. doi:10.1080/14772019.2016.1208687. 
  7. ↑ Mothé, Dimila; Ferretti, Marco P.; Avilla, Leonardo S. (12 January 2016). "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. doi:10.1371/journal.pone.0147009. PMID 26756209. Bibcode: 2016PLoSO..1147009M. 
  8. ↑ Lambert, W. David (2023-10-02). "Implications of discoveries of the shovel-tusked gomphothere Konobelodon (Proboscidea, Gomphotheriidae) in Eurasia for the status of Amebelodon with a new genus of shovel-tusked gomphothere, Stenobelodon" (in en). Journal of Vertebrate Paleontology. doi:10.1080/02724634.2023.2252021. ISSN 0272-4634. https://www.tandfonline.com/doi/full/10.1080/02724634.2023.2252021. 
  9. ↑ Wu, Yan; Deng, Tao; Hu, Yaowu; Ma, Jiao; Zhou, Xinying; Mao, Limi; Zhang, Hanwen; Ye, Jie et al. (2018-05-16). "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. 
  10. ↑ Smith, Gregory James; DeSantis, Larisa R. G. (February 2020). "Extinction of North American Cuvieronius (Mammalia: Proboscidea: Gomphotheriidae) driven by dietary resource competition with sympatric mammoths and mastodons". Paleobiology 46 (1): 41–57. doi:10.1017/pab.2020.7. ISSN 0094-8373. Bibcode: 2020Pbio...46...41S. 
  11. ↑ El Adli, Joseph J.; Fisher, Daniel C.; Cherney, Michael D.; Labarca, Rafael; Lacombat, Frédéric (July 2017). "First analysis of life history and season of death of a South American gomphothere" (in en). Quaternary International 443: 180–188. doi:10.1016/j.quaint.2017.03.016. Bibcode: 2017QuInt.443..180E. https://linkinghub.elsevier.com/retrieve/pii/S1040618216302634. 
  12. ↑ Mothé, Dimila; Avilla, Leonardo S.; Winck, Gisele R. (December 2010). "Population structure of the gomphothere Stegomastodon waringi (Mammalia: Proboscidea: Gomphotheriidae) from the Pleistocene of Brazil" (in en). Anais da Academia Brasileira de Ciências 82 (4): 983–996. doi:10.1590/S0001-37652010005000001. ISSN 0001-3765. PMID 21152772. http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0001-37652010000400020&lng=en&nrm=iso&tlng=en. 
  13. ↑ Sanders, William J. (2023-07-07) (in en). Evolution and Fossil Record of African Proboscidea (1 ed.). Boca Raton: CRC Press. pp. 94. doi:10.1201/b20016. ISBN 978-1-315-11891-8. https://www.taylorfrancis.com/books/9781315118918. 
  14. ↑ 14.0 14.1 Cantalapiedra, Juan L.; Sanisdro, Oscar L.; Zhang, Hanwen; Alberdi, Mª Teresa; Prado, Jose Luis; Blanco, Fernando; Saarinen, Juha (1 July 2021). "The rise and fall of proboscidean ecological diversity". Nature Ecology & Evolution 355 (9): 1266–1272. doi:10.1038/s41559-021-01498-w. PMID 34211141. https://www.nature.com/articles/s41559-021-01498-w. Retrieved 21 August 2021. 
  15. ↑ 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): e1318284. doi:10.1080/02724634.2017.1318284. ISSN 0272-4634. Bibcode: 2017JVPal..37E8284W. https://www.tandfonline.com/doi/full/10.1080/02724634.2017.1318284. 
  16. ↑ Spencer LG 2022. The last North American gomphotheres. N Mex Mus Nat Hist Sci. 88:45–58.
  17. ↑ Wang, Shi-Qi; Saegusa, Haruo; Duangkrayom, Jaroon; He, Wen; Chen, Shan-Qin (December 2017). "A new species of Tetralophodon from the Linxia Basin and the biostratigraphic significance of tetralophodont gomphotheres from the Upper Miocene of northern China" (in en). Palaeoworld 26 (4): 703–717. doi:10.1016/j.palwor.2017.03.005. https://linkinghub.elsevier.com/retrieve/pii/S1871174X16301366. 
  18. ↑ 18.0 18.1 Parray, Khursheed A.; Jukar, Advait M.; Paul, Abdul Qayoom; Ahmad, Ishfaq; Patnaik, Rajeev (March 2022). Silcox, Mary. ed. "A gomphothere (Mammalia, Proboscidea) from the Quaternary of the Kashmir Valley, India" (in en). Papers in Palaeontology 8 (2). doi:10.1002/spp2.1427. ISSN 2056-2799. https://onlinelibrary.wiley.com/doi/10.1002/spp2.1427. 
  19. ↑ Konidaris, George E.; Tsoukala, Evangelia (2022), Vlachos, Evangelos, ed., "The Fossil Record of the Neogene Proboscidea (Mammalia) in Greece" (in en), Fossil Vertebrates of Greece Vol. 1 (Cham: Springer International Publishing): pp. 299–344, doi:10.1007/978-3-030-68398-6_12, ISBN 978-3-030-68397-9, https://link.springer.com/10.1007/978-3-030-68398-6_12, retrieved 2023-03-25 
  20. ↑ Sanders, William J.; Haile-Selassie, Yohannes (June 2012). "A New Assemblage of Mid-Pliocene Proboscideans from the Woranso-Mille Area, Afar Region, Ethiopia: Taxonomic, Evolutionary, and Paleoecological Considerations" (in en). Journal of Mammalian Evolution 19 (2): 105–128. doi:10.1007/s10914-011-9181-y. ISSN 1064-7554. http://link.springer.com/10.1007/s10914-011-9181-y. 
  21. ↑ 21.0 21.1 Mothé, Dimila; dos Santos Avilla, Leonardo; Asevedo, Lidiane; Borges-Silva, Leon; Rosas, Mariane; Labarca-Encina, Rafael; Souberlich, Ricardo; Soibelzon, Esteban et al. (30 September 2016). "Sixty years after 'The mastodonts of Brazil': The state of the art of South American proboscideans (Proboscidea, Gomphotheriidae)". Quaternary International 443: 52–64. doi:10.1016/j.quaint.2016.08.028. Bibcode: 2017QuInt.443...52M. http://bibdigital.epn.edu.ec/bitstream/15000/17075/1/Moth%c3%a9%20et%20al.%2c%202016%20-%20Sixty%20years%20proboscideans.pdf. 
  22. ↑ Konidaris, George E.; Tsoukala, Evangelia (2022), Vlachos, Evangelos, ed., "The Fossil Record of the Neogene Proboscidea (Mammalia) in Greece" (in en), Fossil Vertebrates of Greece Vol. 1 (Cham: Springer International Publishing): pp. 299–344, doi:10.1007/978-3-030-68398-6_12, ISBN 978-3-030-68397-9, https://link.springer.com/10.1007/978-3-030-68398-6_12, retrieved 2023-03-23 
  23. ↑ Konidaris, George E.; Roussiakis, Socrates J. (2018-11-02). "The first record of Anancus (Mammalia, Proboscidea) in the late Miocene of Greece and reappraisal of the primitive anancines from Europe" (in en). Journal of Vertebrate Paleontology 38 (6): e1534118. doi:10.1080/02724634.2018.1534118. ISSN 0272-4634. Bibcode: 2018JVPal..38E4118K. https://www.tandfonline.com/doi/full/10.1080/02724634.2018.1534118. 
  24. ↑ Rivals, Florent; Mol, Dick; Lacombat, Frédéric; Lister, Adrian M.; Semprebon, Gina M. (August 2015). "Resource partitioning and niche separation between mammoths (Mammuthus rumanus and Mammuthus meridionalis) and gomphotheres (Anancus arvernensis) in the Early Pleistocene of Europe" (in en). Quaternary International 379: 164–170. doi:10.1016/j.quaint.2014.12.031. Bibcode: 2015QuInt.379..164R. https://linkinghub.elsevier.com/retrieve/pii/S1040618214009860. 
  25. ↑ Wang, Yuan; Jin, Chang-zhu; Mead, Jim I. (August 2014). "New remains of Sinomastodon yangziensis (Proboscidea, Gomphotheriidae) from Sanhe karst Cave, with discussion on the evolution of Pleistocene Sinomastodon in South China" (in en). Quaternary International 339-340: 90–96. doi:10.1016/j.quaint.2013.03.006. Bibcode: 2014QuInt.339...90W. https://linkinghub.elsevier.com/retrieve/pii/S1040618213001390. 
  26. ↑ 26.0 26.1 Lister, Adrian M. (August 2013). "The role of behaviour in adaptive morphological evolution of African proboscideans" (in en). Nature 500 (7462): 331–334. doi:10.1038/nature12275. ISSN 0028-0836. PMID 23803767. Bibcode: 2013Natur.500..331L. http://www.nature.com/articles/nature12275. 

External links

Wikidata ☰ Q387585 entry

de:Gomphotherium



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