Gomphothere

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Gomphothere
Temporal range: 28–0.01  Ma
Gophotherium at AMNH.jpg
Specimen of Gomphotherium productum at the American Museum of Natural History
Stegomastodon CCB.JPG
Notiomastodon platensis Centro Cultural del Bicentenario de Santiago del Estero in Argentina
Scientific classification OOjs UI icon edit-ltr.svg
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. 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 as part of the Great American Interchange. Gomphotheres are a paraphyletic group ancestral to Elephantidae, which contains modern elephants, as well as Stegodontidae.

Contents

While most famous forms such as Gomphotherium had long lower jaws with tusks, the ancestral condition for the group, some later members developed shortened (brevirostrine) lower jaws with either vestigial or no lower tusks and outlasted the long-jawed gomphotheres. This change made them look very similar to modern elephants, an example of parallel evolution. During the Pliocene and Early Pleistocene, the diversity of gomphotheres declined, ultimately becoming extinct outside of the Americas. The last two genera, Cuvieronius ranging southern North America to western South America, and Notiomastodon ranging over most of South America, continued to exist until the end of the Pleistocene around 12,000 years ago, when they became extinct along with many other megafauna species 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 Knight Gomphotherium.jpg
Life restoration of Gomphotherium
Life restoration of Cuvieronius, a brevirostrine gomphothere Cuvieronius hyodon Mexico.png
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 rather than sharp 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] with this oblique movement being combined with vertical (orthal) motion that served to crush food. [3] Like modern elephants and other members of Elephantimorpha, gomphotheres had horizontal tooth replacement, where teeth would progressively migrate towards the front of the jaws before they were taken place by more posterior teeth. Unlike modern elephants, many gomphotheres retained permanent premolar teeth [4] though they were absent in some gomphothere genera. [5]

Early gomphotheres had lower jaws with an elongate (longitostrine) mandibular symphysis (the front-most part of the lower jaw) 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. [5] In Gomphotheriidae, these elongate mandibular symphysis tend to be narrow, while the lower tusks tend to be club shaped. [6] While the musculature of the trunk of longirostrine gomphotheres was likely very similar to that of living elephants, the trunk was likely shorter (probably no longer than the tips of the lower tusks), and rested upon the elongate lower jaw, though the trunks of later brevirostrine gomphotheres were likely free hanging and comparable to those of living elephants in length. [3] The lower tusks and long lower jaws of primitive gomphotheres were likely used for cutting vegetation, with a secondary contribution in acquiring food using the trunk, while brevirostrine gomphotheres relied primarily on their trunks to acquire food similar to modern elephants. [6] The upper tusks of primitive longirostrine gomphotheres typically gently curve downwards, [3] and generally do not exceed 2 metres (6.6 ft) in length and 35 kilograms (77 lb) in weight, [7] though some later brevirostine gomphotheres developed considerably larger upper tusks. [3] [7] Upper tusks of brevirostine gomphotheres include those which are straight [3] and upwardly curved. [8]

Skeletons of Gomphotherium productum (left) and Gomphotherium steinheimense (right) Gomphotherium skeletals.png
Skeletons of Gomphotherium productum (left) and Gomphotherium steinheimense (right)

Most gomphotheres reached sizes equivalent to those of those of the modern Asian elephant (Elephas maximus), though some gomphotheres reached sizes comparable to or somewhat exceeding African bush elephants (Loxodonta africana). [9] The limb bones of gomphotheres like those of mammutids are generally more robust than elephantids, with the legs also tending to be proportionally shorter. Their bodies also tend to be more proportionally elongate than those of living elephants, resulting in gomphotheres being heavier than an elephant at the same shoulder height. [9] [10]

Taxonomy

"Gomphotheres" are assigned to their own family, Gomphotheriidae, but are widely agreed to be a paraphyletic group. The families Choerolophodontidae and Amebelodontidae (the latter of which includes "shovel tuskers" with flattened lower tusks like Platybelodon ) are sometimes considered gomphotheres sensu lato, [11] [12] [13] though some authors argue that Amebelodontidae should be sunk into Gomphotheriidae. [14] 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. [11] Some authors choose to exclude "tetralophodont gomphotheres" from Gomphotheriidae, and instead assign them to the group Elephantoidea. [11] "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. [15]

While the North American long jawed proboscideans Gnathabelodon, Eubelodon and Megabelodon been assigned to Gomphotheriidae in some studies [5] [16] other studies suggest that they should be assigned to Amebelodontidae (Eubelodon, Megabelodon) or Choerolophodontidae (Gnathabelodon). [6]

Cladogram of Elephantimorpha after Li et al. 2023, showing a paraphyletic Gomphotheriidae. [6]

Elephantimorpha

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 climatic conditions and competition. [17] Analysis of the tusks of a male Notiomastodon individual suggest that it underwent musth, similar to modern elephants. [18] Notiomastiodon is also suggested to have lived in social family groups, like modern elephants. [19]

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. [20] Gomphotheres arrived in Eurasia after the connection of Afro-Arabia and Eurasia during the Early Miocene around 19 million years ago, [21] in what is termed the "Proboscidean Datum Event". Gomphotherium arrived in North America around 16 million years ago, [22] and is suggested to be the ancestor of later New World gomphothere genera. [23] "Trilophodont gomphotheres" dramatically declined during the Late Miocene, likely due to the increasing C4 grass-dominated habitats, [21] while during the Late Miocene "tetralophodont gomphotheres" were abundant and widespread in Eurasia, where they represented the dominant group of proboscideans. [24] All trilophodont gomphotheres, with the exception of the Asian Sinomastodon , became extinct in Eurasia by the beginning of the Pliocene, [25] along with the global extinction of the "shovel tusker" amebelodontids. [26] 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. [27] 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, becoming widespread across the continent. [28] The last gomphothere native to Europe, Anancus arvernensis [29] became extinct during the Early Pleistocene, around 1.6–2 million years ago [30] [31] Sinomastodon became extinct at the end of the Early Pleistocene, around 800,000 years ago. [32] From the latter half of the Early Pleistocene onwards, gomphotheres were extirpated from most of North America, likely due to competition with mammoths and mastodons. [17]

The extinction of gomphotheres in Afro-Eurasia has generally been supposed to be the result the expansion of Elephantidae and Stegodon. [25] [33] The morphology of elephantid molars being more efficient than gomphotheres in consuming grass, which became more abundant during the Pliocene and Pleistocene epochs. [33] In southern North America, Central America and South America, gomphotheres did not become extinct until shortly after the arrival of humans to the Americas, approximately 12,000 years ago, as part of the Late Pleistocene megafauna extinctions of most large mammals across the Americas. 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. [28]

Related Research Articles

<span class="mw-page-title-main">Proboscidea</span> Order of mammals including elephants

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 species of elephant are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant.

<span class="mw-page-title-main">Elephantidae</span> Family of mammals

Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are large terrestrial mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the family are extinct. Only two genera, Loxodonta and Elephas, are living.

<span class="mw-page-title-main">Mammutidae</span> Extinct family of mammals

Mammutidae is an extinct family of proboscideans belonging to Elephantimorpha. It is best known for the mastodons, which inhabited North America from the Late Miocene until their extinction at beginning of the Holocene, around 11,000 years ago. The earliest fossils of the group are known from the Late Oligocene of Africa, around 24 million years ago, and fossils of the group have also been found across Eurasia. The name "mastodon" derives from Greek, μαστός "nipple" and ὀδούς "tooth", referring to their characteristic teeth.

<i>Gomphotherium</i> Extinct genus of elephant-like mammals

Gomphotherium is an extinct genus of gomphothere proboscidean from the Neogene 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.

<i>Anancus</i> Genus of proboscideans

Anancus is an extinct genus of "tetralophodont gomphothere" native to Afro-Eurasia, that lived from the Tortonian stage of the late Miocene until its extinction during the Early Pleistocene, roughly from 8.5–2 million years ago.

<i>Cuvieronius</i> Extinct genus of mammals

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 2.3 metres (7.5 ft) and a body mass of 3.5 tonnes (7,700 lb), it was on average shorter but comparable in body mass to an Asian elephant. Cuvieronius inhabited subtropical and tropical latitudes in environments ranging from grasslands to tropical rainforest. Among the last gomphotheres along with the South American Notiomastodon, it became extinct as part of the end Pleistocene-extinction event, approximately 12,000 years ago, along with most other large mammals in the Americas. The extinctions followed the arrival of humans to the Americas, and evidence has been found for human hunting of Cuvieronius, which may have been a factor in its extinction.

<i>Stegomastodon</i> Extinct genus of mammals

Stegomastodon is an extinct genus of gomphotheres. It ranged throughout North America from the Pliocene, to the Early Pleistocene. The former South American species have been synonymized with Notiomastodon platensis.

<i>Sinomastodon</i> Extinct genus of gomphothere proboscidean

Sinomastodon is an extinct gomphothere genus known from the Late Miocene to Early Pleistocene of Asia, including China, Japan, Thailand, Myanmar, Indonesia and probably Kashmir.

<i>Tetralophodon</i> Extinct genus of mammals

Tetralophodon is an extinct genus of "tetralophodont gomphothere" belonging to the superfamily Elephantoidea, known from the Miocene of Afro-Eurasia.

<i>Rhynchotherium</i> Extinct genus of proboscid

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 Ma, living for approximately 10 million years.

Gnathabelodon is an extinct genus of gomphothere endemic to North America that includes species that lived during the Middle to Late Miocene.

<i>Eubelodon</i> Extinct genus of proboscid

Eubelodon is an extinct genus of gomphothere which lived in North America during the Miocene Epoch. It contains a single species: Eubelodon morrilli.

<i>Notiomastodon</i> Extinct genus of gomphothere elephantimorph native to South America

Notiomastodon is an extinct genus of gomphothere proboscidean, endemic to South America from the Pleistocene to the beginning of the Holocene. Notiomastodon specimens reached a size similar to that of the modern Asian elephant, with a body mass of 3-4 tonnes. Like other brevirostrine gomphotheres such as Cuvieronius and Stegomastodon, Notiomastodon had a shortened lower jaw and lacked lower tusks, unlike more primitive gomphotheres like Gomphotherium.

<span class="mw-page-title-main">Stegodontidae</span> Family of extinct elephant-like mammals

Stegodontidae is an extinct family of proboscideans from Africa and Asia from the Early Miocene to the Late Pleistocene. It contains two genera, the earlier Stegolophodon, known from the Miocene of Asia and the later Stegodon, from the Late Miocene to Late Pleistocene of Africa and Asia which is thought to have evolved from the former. The group is noted for their plate-like lophs on their teeth, which are similar to elephants and different from those of other extinct proboscideans like gomphotheres and mammutids, with both groups having a proal jaw movement utilizing forward strokes of the lower jaw. These similarities with modern elephants were probably convergently evolved. Like elephantids, stegodontids are thought to have evolved from gomphothere ancestors.

<i>Choerolophodon</i> Extinct genus of mammals

Choerolophodon is an extinct genus of proboscidean that lived during the Miocene of Eurasia and Africa. Fossils of Choerolophodon have been found in Africa, Southeast Europe, Turkey, Iraq, Iran, the Indian subcontinent, and China.

<span class="mw-page-title-main">Elephantimorpha</span> Clade of mammals

Elephantimorpha is a clade of proboscideans that contains the Mammutidae (mastodons), as well as Elephantida. All members of this group have the horizontal tooth replacement typical of modern elephants, unlike more primitive members of the Elephantiformes. Like modern elephants, the ancestor of Elephantimorpha was likely capable of communicating via infrasonic calls. While early elephantimorphs generally had lower jaws with an elongated mandibular symphysis at the front of the jaw with well developed lower tusks/incisors, from the Late Miocene onwards, many groups convergently developed brevirostrine (shortened) lower jaws with vestigial or no lower tusks, probably corresponding with the elongation and increasingly dexterity of the trunk allowing it to be used as the primary feeding organ.

<i>Konobelodon</i> Extinct genus of mammals

Konobelodon is an extinct genus of amebelodont proboscidean from the Miocene of Africa, Eurasia and North America.

<span class="mw-page-title-main">Amebelodontidae</span> Extinct family of mammals

Amebelodontidae is an extinct family of large herbivorous proboscidean mammals related to elephants. They were formerly assigned to Gomphotheriidae, but recent authors consider them a distinct family. They are distinguished from other proboscideans by having flattened lower tusks and very elongate mandibular symphysis. The lower tusks could grow to considerable size, with those of Konobelodon reaching 1.61 metres (5.3 ft) in length. Their molar teeth are typically trilophodont, and possessed posttrite conules. In the past, amebelodonts' shovel-like mandibular tusks led to them being portrayed scooping up water plants, however, dental microwear suggests that they were browsers and mixed feeders. The lower tusks have been proposed to have had a variety of functions depending on the species, including stripping bark, cutting through vegetation, as well as possibly digging. They first appeared in Africa during the Early Miocene, and subsequently dispersed into Eurasia and then North America. They became extinct by the beginning of the Pliocene. While some phylogenetic studies have recovered Amebelodontidae as a monophyletic group that forms the sister group to Gomphotheriidae proper, some authors have argued that Amebelodontidae may be polyphyletic, with it being suggested that the shovel-tusked condition arose several times independently within Gomphotheriidae, thus rendering the family invalid.

<span class="mw-page-title-main">Choerolophodontidae</span> Extinct family of mammals

Choerolophodontidae is an extinct family of proboscideans belonging to Elephantida. Two genera are widely recognised, Afrochoerodon and Choerolophodon.

Eurybelodon is an extinct genus of proboscidean in the family Amebelodontidae. It lived in the Clarendonian age of the Miocene.

References

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  29. ↑ Konidaris, George E.; Tsoukala, Evangelia (2022), Vlachos, Evangelos (ed.), "The Fossil Record of the Neogene Proboscidea (Mammalia) in Greece", 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, S2CID   245023119 , retrieved 2023-03-23
  30. ↑ 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". Journal of Vertebrate Paleontology. 38 (6): e1534118. Bibcode:2018JVPal..38E4118K. doi:10.1080/02724634.2018.1534118. ISSN   0272-4634. S2CID   91391249.
  31. ↑ 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". Quaternary International. 379: 164–170. Bibcode:2015QuInt.379..164R. doi:10.1016/j.quaint.2014.12.031.
  32. ↑ 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". Quaternary International. 339–340: 90–96. Bibcode:2014QuInt.339...90W. doi:10.1016/j.quaint.2013.03.006.
  33. 1 2 Lister, Adrian M. (August 2013). "The role of behaviour in adaptive morphological evolution of African proboscideans". Nature. 500 (7462): 331–334. Bibcode:2013Natur.500..331L. doi:10.1038/nature12275. ISSN   0028-0836. PMID   23803767. S2CID   883007.
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