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Introduction / Physical Description

Gomphotheres were early elephant relatives that looked different from modern elephants. Most species had four tusks: two straight tusks in the upper jaw and two straight tusks in the lower jaw. Unlike modern elephant tusks, the upper tusks of many gomphotheres were coated with enamel. Their ridged molars were ideal for grinding tough foods such as branches, bark, and shrubs.

The skull of Gomphotherium was elongated, and the body was more barrel-shaped than that of modern elephants. Larramendi (2016) examined the body proportions of several proboscideans and found that most gomphotheres had very short tails. Although they are often shown with little hair, Larramendi also suggests they may have had a covering of hair. Their trunks were probably long enough to reach the ground without bending their knees. Most species were similar in size to the Asian elephant, although G. steinheimense was larger.

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Origins, Distribution, and Species

Gomphotheres were a very successful group of early elephant relatives. They are associated with a major proboscidean radiation in which elephants and their relatives migrated out of Africa and spread into Europe, Asia, and the Americas. Although the genus Gomphotherium is extinct, G. steinheimense is considered part of the evolutionary line that gave rise to modern elephants (Wu Y. et al., 2018).

Origin and Distribution: Gomphothere-like animals first appear in the fossil record in the late Oligocene of eastern Africa. A notable example is Eritreum melakeghebrekristosi from Eritrea, near Ethiopia (Shoshani, 2006). By the early Miocene, gomphotheres were present in Africa, Europe, Asia, and Japan (Miyata and Tomida, 2010). By the Middle Miocene, they had spread into North America, and by the Late Miocene to Pliocene they were also found in Central America. The southernmost occurrence of the genus Gomphotherium is from Panama (MacFadden et al., 2015). Close relatives in the Gomphotheriidae family eventually spread into South America during the Pliocene and Pleistocene.

Species: Like many fossil taxa, the Gomphotherium taxonomy is complicated and probably needs revision. Up to 20 species have been accepted at various times, although many are based on isolated material. To simplify the group, Gomphotherium is often divided into four evolutionary groups with a range of species in each group.

1. The Old World Group: This ancestral group is often called the annectens group. It contains earlier species from Africa, Europe, and Asia, including G. annectens, G. sylvaticum, and G. cooperi.

2. The African Taxa: This group is a separate evolutionary line unique to Africa. It includes G. libycum and G. pygmaeus.

3. The Angustidens Group: This is another separate evolutionary group in Europe and Asia. It includes G. inopinatum, G. mongoliense, G. connexum, G. angustidens, and the Chinese species G. shensiensis.

4. The Derived Gomphotherium Group: This group contains later gomphotheres from the Middle to Late Miocene in Europe, Asia, and North America. It includes G. subtapiroideum, G. tassyi, G. wimani, G. browni, and G. steinheimense.

North and Central American species are also included in the derived group. American gomphothere species include G. productum from the Midwest, G. calvertensis from the Calvert Cliffs of Maryland and Virginia, G. cimarronis from Texas, G. simplicidens from Florida, and G. hondurensis from Central America.

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Ecology and Diet: Woodland Browsers

Gomphotheres had molars with high ridges, making them efficient at grinding tough vegetation such as tree limbs, bark, and shrubs. Most gomphotheres probably lived in wooded environments and used their tusks to shear vegetation from trees and shrubs. A common misconception is that gomphotheres lived in swampy environments and used their shovel-like lower tusks to scoop vegetation from lakes, but this is not supported by wear and isotope studies.

Fox and Fisher (2004) studied carbon isotope ratios in tusks to reconstruct the dietary habits of North American gomphotheres and found that they stayed in wooded habitats and fed on trees and shrubs. Similar results were found in China by Zhang et al. (2016). Separate studies of tusk wear patterns also suggest that gomphotheres used their upper and lower tusks to shear bark and vegetation from trees rather than scoop aquatic plants. An extreme example is Platybelodon, a related gomphothere with wide, flattened lower tusks.

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Extinction: The Expansion of Grasslands

Gomphotheres were early elephant relatives well suited for eating tough browse such as branches and shrubs. As the Miocene came to an end, the global climate became cooler and more arid (Chamberlain et al., 2014). Forested environments gave way to grasslands, influencing mammal evolution around the world (Cerling et al., 1993; Janis, 1993).

Although climates and ecosystems changed, the diet of Gomphotherium appears to have remained largely browser-based. Research indicates that North American gomphotheres continued feeding in wooded environments (Fox and Fisher, 2004), and similar patterns are seen in China (Zhang et al., 2016). As specialized grazing mammals spread and flourished, traditional gomphothere habitats declined.

By the middle Pliocene, the genus Gomphotherium had disappeared. It was replaced by more modern elephants with flatter-ridged molars better suited for grinding grasses. Although Gomphotherium went extinct, a few members of the Gomphotheriidae family survived into the Ice Ages. Examples include Cuvieronius and Notiomastodon, which migrated into South America. These later gomphotheres had reduced lower tusks, lost enamel on the upper tusks, and developed mixed browsing and grazing diets. They disappeared relatively recently, around the end of the Pleistocene and beginning of the Holocene.

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Works Cited / References

Cerling, T., Wang, Y., and Quade, J. (1993). Expansion of C4 ecosystems as an indicator of global ecological change in the late Miocene. Nature, 361, 344-345.

Chamberlain, C., Winnick, M., Hari, M., Chamberlain, S., and Katharine M. (2014). The impact of Neogene grassland expansion and aridification on the isotopic composition of continental precipitation. Global Biogeochemical Cycles. DOI: 10.1002/2014GB004822.

Fox, D., and Fisher, D. (2004). Dietary reconstruction of Miocene Gomphotherium (Mammalia, Proboscidea) from the Great Plains region, USA, based on the carbon isotope composition of tusk and molar enamel. Palaeogeography, Palaeoclimatology, Palaeoecology, 206, 311-335. DOI: 10.1016/j.palaeo.2004.01.010.

Janis, C. M. (1993). Mammal evolution in the context of changing climates, vegetation and tectonic events. Annual Review of Ecology and Systematics, 24, 467-500.

Lambert, W. (1992). The feeding habits of the shovel-tusked gomphotheres: evidence from tusk wear patterns. Paleobiology, 18.

Larramendi, A. (2015). Shoulder Height, Body Mass and Shape of Proboscideans. Acta Palaeontologica Polonica, 61. DOI: 10.4202/app.00136.2014.

Lucas, S. and Morgan, G. (2005). Ice Age Proboscideans of New Mexico. New Mexico Museum of Natural History and Science Bulletin, 28, 255-261.

MacFadden, B., Morgan, G., Jones, D., and Rincon, A. (2015). Gomphothere proboscidean (Gomphotherium) from the late Neogene of Panama. Journal of Paleontology, 89(2), 360-365. DOI: 10.1017/jpa.2014.31.

Mothe, D., Ferretti, M. P., and Avilla, L. S. (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.

Mothe, D., dos Santos Avilla, L., Araújo Júnior, H., Rotti, A., Prous, A., and Azevedo, S.A.K. (2020). An artifact embedded in an extinct proboscidean sheds new light on human-megafaunal interactions in the Quaternary of South America. Quaternary Science Reviews, 229, 106125. DOI: 10.1016/j.quascirev.2019.106125.

Prado, J.L. and Alberdi, M.T. (2008). A cladistic analysis among trilophodont gomphotheres (Mammalia, Proboscidea) with special attention to the South American genera. Palaeontology, 51, 903-915. DOI: 10.1111/j.1475-4983.2008.00785.x.

Shoshani, J., Walter, R. C., Abraha, M., Berhe, S., Tassy, P., Sanders, W. J., Marchant, G. H., Libsekal, Y., Ghirmai, T., and Zinner, D. (2006). A proboscidean from the late Oligocene of Eritrea, a missing link between early Elephantiformes and Elephantimorpha, and biogeographic implications. Proceedings of the National Academy of Sciences, 103(46), 17296-17301. DOI: 10.1073/pnas.0603689103.

Pérez-Crespo, V. A., Prado, J. L., Alberdi, M. T., Arroyo-Cabrales, J., and Johnson, E. (2016). Diet and Habitat for Six American Pleistocene Proboscidean Species Using Carbon and Oxygen Stable Isotopes. Ameghiniana, 53(1), 39-51. DOI: 10.5710/AMGH.02.06.2015.2842.

Miyata, K. and Tomida, Y. (2010). Anchitherium from the early Miocene Hiramaki Formation, Gifu Prefecture, Japan, and its implication for the early diversification of Asian Anchitherium. Journal of Paleontology, 84, 763-773. DOI: 10.1017/S0022336000058479.

Wang, Shiqi, Li, Yu, Duangkrayom, Jaroon, Yang, Xiang-Wen, He, Wen, and Chen, Shan-Qin. (2017). A new species of Gomphotherium (Proboscidea, Mammalia) from China and the evolution of Gomphotherium in Eurasia. Journal of Vertebrate Paleontology, e1318284. DOI: 10.1080/02724634.2017.1318284.

Wu, Yan, Deng, Tao, Hu, Yaowu, Ma, Jiao, Zhou, Xinying, Mao, Limi, Zhang, Hanwen, Ye, Jie, and Wang, Shi-Qi. (2018). A grazing Gomphotherium in Middle Miocene Central Asia, 10 million years prior to the origin of the Elephantidae. Scientific Reports, 8(1). DOI: 10.1038/s41598-018-25909-4.

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Additional Images of Gomphotherium

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