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Amynodontidae

Amynodontidae is an extinct family of perissodactyls related to modern rhinoceroses, known from the Eocene to Oligocene of North America, Europe, and Asia. They were among the dominant groups of herbivorous mammals in the Northern Hemisphere, and were most diverse in Asia. Amynodonts have been nicknamed "hippo-like rhinos" due to their hippopotamus-like build, with massive bodies and short, robust limbs, as well as their tusk-like canine teeth.

Description
Amynodonts have been nicknamed "hippo-like rhinos" since they had several hippopotamus-like features, such as massive bodies and short, robust limbs. They were medium-sized to large animals. In the more derived amynodont groups, the condition of the fossae becomes a distinguishing trait; metamynodontines have reduced fossae whereas cadurcodontines have fossae that extend medial to the orbit (towards the middle of the orbits). This further differentiates them from other rhinocerotoids, who either have small canines or lack them entirely. When other rhinocerotoid groups developed tusks, they were typically developed from the incisor teeth. == History of discovery ==
History of discovery
The first amynodont to be described was Cadurcotherium cayluxi in 1873, described by Paul Gervais from the Oligocene of France. The first known North American amynodont fossils were found in the Uinta Formation of Utah and described by Othniel Charles Marsh. In 1875, Marsh named the new species Diceratherium advenum based on a fossil skull. Marsh soon realized that the fossil was distinct from Diceratherium, a previously described true rhinoceros (Rhinocerotidae), and moved it to the new genus Amynodon, as A. advenus, in 1877. In 1887, the new amynodont genus Metamynodon was described based on fossils from the White River Group. Based on Metamynodon, William Berryman Scott and Henry Fairfield Osborn were able to establish several important family-level characteristics of the amynodonts, including the enlarged sagittal crest and the loss of P1 (the first premolar tooth). The first discoveries of amynodonts in Asia were made in Oligocene deposits in Pakistan and in Eocene deposits in Myanmar in the 1910s and 1920s. In the 1920s and 1930s, amynodont fossils were also discovered in Mongolia and China. == Classification ==
Classification
External relations (A), the hyracodont Hyracodon (B), the paracerathere Paraceratherium (C), and the rhinoceros Trigonias'' (D)|upright=1.15 Marsh believed Amynodon to be a rhinoceros and did not separate it from other rhinoceroses above the genus level. In 1889, Osborn published the first detailed comparison of Amynodontidae, Hyracodontidae, and Rhinocerotidae, and established the Amynodontidae as clearly distinct based mainly on their large canines and their shortened face (a trait later found to only apply to derived members of the family). Internal systematics '', from Late Eocene or Early Oligocene Romania The first attempt to establish the relations between amynodont genera was done by Miklós Kretzoi in 1942. Kretzoi established four amynodont subfamilies: Amynodontinae, Cadurcotheriinae, Metamynodontinae, and Paramynodontinae. In 2017, Alexander Averianov, Igor Danilov, Jin Jianhua , and Wang Yingyong published a new phylogenetic analysis of the amynodonts as part of their description of the species Cadurcodon maomingensis. Averianov et al. considered Procadurcodon to be valid, with Z. protheroi as a possible synonym, whereas Tissier et al. considered Procadurcodon dubious and Z. protheroi valid. In 2023, a more comprehensive phylogenetic analysis by Léa Veine-Tonizzo, Tissier, Maia Bukhisianidze, Davit Vasilyan, and Damien Becker produced a result similar to the 2018 analysis. Topology A: Averianov et al. (2017) Topology B: Tissier et al. (2018) Topology C: Veine-Tonizzo et al. (2023) == List of genera ==
List of genera
Additional amynodont fossils have been reported but their taxonomic attribution at the genus level is questionable. These include "Amynodon" hungaricus from the Late Eocene of Hungary, "Amynodon" sinensis from the Late Eocene of Henan, China, "Amynodon" watanabei from the Middle–Late Eocene of Japan, and the gigantic "Metamynodon" bugtiensis from the Oligocene of Pakistan. Revisions have variously retained "M". bugtiensis as an amynodont or referred it to the paracerathere Paraceratherium. == Evolutionary history and paleobiogeography ==
Evolutionary history and paleobiogeography
'', an early amynodont known from the Early to Middle Eocene of Inner Mongolia in China Amynodonts probably originated in Asia during the Early Eocene. The oldest known Asian amynodont is Andarakodon, from the Early Eocene of Kyrgyzstan. The Early–Middle Eocene deposits of the Irdin Manha Formation in China have yielded fossils of the oldest well-defined member of the group, Rostriamynodon. Caenolophus, contemporary with Rostriamynodon, has been suggested to be the most primitive amynodont but its placement in Amynodontidae is disputed; Caenolophus has also been classified as a hyracodont, and appears to be anatomically intermediate between the two families. Teilhardia, known from the same area, has also been suggested to be either a hyracodont or a primitive amynodont but it is fragmentarily known and possibly synonymous with Caenolophus. Amynodonts spread to North America shortly after their initial appearance and achieved a Holarctic distribution. They became one of the dominant herbivore groups of their time. In North America, the temporal range of the amynodonts extends from the Middle Eocene to the Early Oligocene. The earliest known North American example of the group is Amynodon, from the late Bridgerian and Uintan land mammal ages. The amynodonts were at their greatest taxonomic diversity in Asia, especially in Central Asia. Amynodonts increased in body size over the course of their evolutionary history, the smallest known amynodont is the relatively basal "Amynodon" sinensis, estimated at 127 ± 15 kg (280 ± 33 lbs), whereas the largest are the derived Zaisanamynodon borisovi, at 2442 ± 257 kg (5384 ± 567 lbs), and Procadurcodon orientalis, at 2720 kg (5997 lbs).) of Cadurcotherium nouleti, known from the Oligocene of France and SwitzerlandMigration across Beringia was highly important in amynodont evolution. The ancestors of the North American Amynodon and Amynodontopsis bodei were probably amynodonts that migrated from Asia separately; Amynodontopsis bodei is more derived than Amynodon but likely descended from the Asian species Amynodontopsis jiyuanensis. Phylogenetic analyses suggest that there may have been as many as four or five dispersal events where new amynodonts spread from Asia into North America. There were also several dispersal events into Europe. Cadurcotherium is the only genus recorded from Western Europe but Amynodontopsis, Cadurcodon, and Sellamynodon are all known from Eastern Europe. Amynodonts do not appear to have reached Northern Europe. The migration of amynodonts into Europe in the Oligocene may have been facilitated by the preceding Grande Coupure extinction event. The Middle to Late Eocene was a time of great environmental change, which may have facilitated the evolutionary radiation of the amynodonts. In North America, the environment changed during this time from lush semitropical forests to being dominated by open forests and grasslands. The timeframe corresponding to the late Uintan and the Duchesnean land mammal ages in North America saw the emergence of the cadurcodontines and metamynodontines, illustrating diversification and pursuits of different lifestyles. Similar to North American examples, Asian amynodonts also appear to diversify around the boundary between the Eocene and Oligocene. The metamynodontines experienced evolutionary radiation in North America, represented by the genera Megalamynodon and Metamynodon. The record of carcudontines in North America is sparse and depends on the phylogenetic position of Amynodontopsis and the assessments of the species Zaisanamynodon protheroi and Amynodon reedi. Amynodonts were more successful in Asia than in North America, perhaps due to the increasing aridity in North America going into the Oligocene. Metamynodontines survived in North America into the Oligocene; Metamynodon lived in river system across most of the continent, but was a rare element of the Oligocene fauna and was extinct by the end of the Whitneyan land mammal age (late Early Oligocene). Amynodonts in Europe disappeared in the latest Oligocene. Amynodonts survived the longest in Asia, where Cadurcotherium persisted in modern-day Pakistan until the latest Oligocene. The last Asian amynodonts were previously believed to have reached into the early Miocene, but this dating has proven incorrect. ==References==
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