Ungulates are members of the diverse clade Euungulata, which primarily consists of large mammals with hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined to be a polyphyletic and thereby invalid clade based on molecular data. As a result, true ungulates had since been reclassified to the newer clade Euungulata in 2001 within the clade Laurasiatheria while Paenungulata has been reclassified to a distant clade Afrotheria. Living ungulates are divided into two orders: Perissodactyla including equines, rhinoceroses, and tapirs; and Artiodactyla including cattle, antelope, pigs, giraffes, camels, sheep, deer, and hippopotamuses, among others. Cetaceans such as whales, dolphins, and porpoises are also classified as artiodactyls, although they do not have hooves. Most terrestrial ungulates use the hoofed tips of their toes to support their body weight while standing or moving. Two other orders of ungulates, Notoungulata and Litopterna, both native to South America, became extinct at the end of the Pleistocene, around 12,000 years ago.
Artiodactyls are placental mammals belonging to the order Artiodactyla. Typically, they are ungulates which bear weight equally on two of their five toes. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of the five toes. Another difference between the two orders is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine. Molecular biology, along with new fossil discoveries, has found that cetaceans fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use "even-toed ungulates" to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature.
Andrewsarchus, meaning "Andrews' ruler", is an extinct genus of artiodactyl that lived during the Middle Eocene in what is now China. The genus was first described by Henry Fairfield Osborn in 1924 with the type species A. mongoliensis based on a largely complete cranium. A second species, A. crassum, was described in 1977 based on teeth. A mandible, formerly described as Paratriisodon, does probably belong to Andrewsarchus as well. The genus has been historically placed in the families Mesonychidae or Arctocyonidae, or was considered to be a close relative of whales. It is now regarded as the sole member of its own family, Andrewsarchidae, and may have been related to entelodonts. Fossils of Andrewsarchus have been recovered from the Middle Eocene Irdin Manha, Lushi, and Dongjun Formations of Inner Mongolia, each dated to the Irdinmanhan Asian land mammal age.
Creodonta is a former order of extinct carnivorous placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ancestral to the modern Carnivora, this order is now usually considered a polyphyletic assemblage of two different groups, the oxyaenids and the hyaenodontids, not a natural group. Oxyaenids are first known from the Palaeocene of North America, while hyaenodonts hail from the Palaeocene of Africa.
Entelodontidae is an extinct family of pig-like artiodactyls which inhabited the Northern Hemisphere from the late Eocene to the early Miocene epochs, about 38-19 million years ago. Their large heads, low snouts, narrow gait, and proposed omnivorous diet inspires comparisons to suids and tayassuids (peccaries), and historically they have been considered closely related to these families purely on a morphological basis. However, studies which combine morphological and molecular (genetic) data on artiodactyls instead suggest that entelodonts are cetancodontamorphs, more closely related to hippos and cetaceans through their resemblance to Pakicetus, than to basal pigs like Kubanochoerus and other ungulates.
Mesonychia is an extinct taxon of small- to large-sized carnivorous ungulates related to artiodactyls. Mesonychians first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely when the last genus, Mongolestes, became extinct in the early Oligocene. In Asia, the record of their history suggests they grew gradually larger and more predatory over time, then shifted to scavenging and bone-crushing lifestyles before the group became extinct.
Palaeotherium is an extinct genus of equoid that lived in Europe and possibly the Middle East from the Middle Eocene to the Early Oligocene. It is the type genus of the Palaeotheriidae, a group exclusive to the Palaeogene that was closest in relation to the Equidae, which contains horses plus their closest relatives and ancestors. Fossils of Palaeotherium were first described in 1782 by the French naturalist Robert de Lamanon and then closely studied by another French naturalist, Georges Cuvier, after 1798. Cuvier erected the genus in 1804 and recognized multiple species based on overall fossil sizes and forms. As one of the first fossil genera to be recognized with official taxonomic authority, it is recognized as an important milestone within the field of palaeontology. The research by early naturalists on Palaeotherium contributed to the developing ideas of evolution, extinction, and succession and demonstrating the morphological diversity of different species within one genus.
Merycoidodontoidea, previously known as "oreodonts" or "ruminating hogs," are an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the better known forms were generally hog-like, and the group has traditionally been placed within the Suina, though some recent work suggests they may have been more closely related to camels. "Oreodont" means "mountain teeth," referring to the appearance of the molars. Most oreodonts were sheep-sized, though some genera grew to the size of cattle. They were heavy-bodied, with short four-toed hooves and comparatively long tails.
Patriofelis is an extinct genus of carnivorous placental mammals from the extinct subfamily Oxyaeninae within the extinct family Oxyaenidae. It was a large cat-like predator which lived in North America during the Bridgerian NALMA. Fossils have been found in Wyoming, Colorado, and Oregon.
Mesonychidae is an extinct family of small to large-sized omnivorous-carnivorous mammals. They were endemic to North America and Eurasia during the Early Paleocene to the Early Oligocene, and were the earliest group of large carnivorous mammals in Asia. Once considered a sister-taxon to artiodactyls, recent evidence now suggests no close connection to any living mammal. Mesonychid taxonomy has long been disputed and they have captured popular imagination as "wolves on hooves", animals that combine features of both ungulates and carnivores. Skulls and teeth have similar features to early whales, and the family was long thought to be the ancestors of cetaceans. Recent fossil discoveries have overturned this idea; the consensus is that whales are highly derived artiodactyls. Some researchers now consider the family a sister group either to whales or to artiodactyls, close relatives rather than direct ancestors. Other studies define Mesonychia as basal to all ungulates, occupying a position between Perissodactyla and Ferae. In this case, the resemblances to early whales would be due to convergent evolution among ungulate-like herbivores that developed adaptations related to hunting or eating meat.
Tillodontia is an extinct suborder of eutherian mammals known from the Early Paleocene to Late Eocene of China, the Late Paleocene to Middle Eocene of North America where they display their maximum species diversity, the Middle Eocene of Pakistan, and the Early Eocene of Europe. Leaving no descendants, they are most closely related to the pantodonts, another extinct group. The tillodonts were medium- to large-sized animals that probably fed on roots and tubers in temperate to subtropical habitats.
Phenacodontidae is an extinct family of large herbivorous mammals traditionally placed in the “wastebasket taxon” Condylarthra, which may instead represent early-stage perissodactyls. They lived from the late early Paleocene to early middle Eocene and their fossil remains have been found in North America and Europe. The only unequivocal Asian phenacodontid is Lophocion asiaticus.
Duerotherium is an extinct genus of Palaeogene artiodactyls known only from the Iberian Peninsula during the Middle Eocene, which contains one species D. sudrei. It, like other members of the Anoplotheriidae, was endemic to Western Europe. The anoplotheriine was described from a left fragment of a maxilla from the Mazaterón Formation of the Duero Basin in 2009. Its dentition is mostly typical of the Anoplotheriinae but differs by an elongated plus triangular 3rd upper premolar and very specific traits of the molars. It is thought to have been part of an endemic faunal assemblage that evolved within the Iberian Peninsula by the Middle Eocene, where climates were subtropical.
Cainotheriidae is an extinct family of artiodactyls known from the Late Eocene to Middle Miocene of Europe. They are mostly found preserved in karstic deposits.
Dichodon is an extinct genus of Palaeogene artiodactyls belonging to the family Xiphodontidae. It was endemic to Western Europe and lived from the middle Eocene up to the earliest Oligocene. The genus was first erected by the British naturalist Richard Owen in 1848 based on dental remains from the fossil beds in Hordle, England. He noticed similar dentitions to contemporary artiodactyls like those of the Anoplotheriidae and Dichobunidae and references the name of the genus Dichobune. Eventually, it was found to be more closely related to Xiphodon and now includes 11 species, although one of them may be synonymous.
Pseudhyrax is an extinct genus of archaeohyracid notoungulate. It lived from the Late Eocene to the Early Oligocene, of what is now South America.
Santiagorothia is an extinct genus of interatheriid notoungulate. It lived during the Early Oligocene, and its fossils were discovered in Argentina and Chile.
Periptychus is an extinct genus of mammal belonging to the family Periptychidae. It lived from the Early to Late Paleocene and its fossil remains have been found in North America.
Diplobune is an extinct genus of Palaeogene artiodactyls belonging to the family Anoplotheriidae. It was endemic to Europe and lived from the late Eocene to the early Oligocene. The genus was first erected as a subgenus of Dichobune by Ludwig Rütimeyer in 1862 based on his hypothesis of the taxon being a transitional form between "Anoplotherium" secundaria, previously erected by Georges Cuvier in 1822, and Dichobune. He based the genus etymology off of the two-pointed pillarlike shapes of the lower molars, which had since been a diagnosis of it. However, in 1870, Diplobune was elevated to genus rank by Oscar Fraas, who recognized that Diplobune was a distinct genus related to Anoplotherium and not Dichobune. After several revisions of the anoplotheriids, there are currently four known species of which D. minor is the type species.
Catodontherium is an extinct genus of Palaeogene artiodactyls belonging to the family Anoplotheriidae. It was endemic to Western Europe and had a temporal range exclusive to the middle Eocene, although its earliest appearance depends on whether C. argentonicum is truly a species of Catodontherium. It was first named Catodus by the French palaeontologist Charles Depéret in 1906, who created two species for the genus and later changed the genus name to Catodontherium in 1908. The Swiss palaeontologist Hans Georg Stehlin renamed one species and classified two other newly erected species to Catodontherium in 1910. Today, there are four known species, although two remain questionable in genus placement.
