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 third and fourth, often in the form of a hoof. 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 is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine as perissodactyls do. The advent of molecular biology, along with new fossil discoveries, 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.
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.
Diacodexis is an extinct genus of small herbivorous mammals belonging to the family Diacodexeidae that lived in North America, Europe and Pakistan from 55.4 mya to 46.2 mya and existing for approximately 9.2 million years.
Elomeryx is an extinct genus of artiodactyl ungulate, and is among the earliest known anthracotheres. The genus was extremely widespread, first being found in Asia in the middle Eocene, in Europe during the latest Eocene, and having spread to North America by the early Oligocene. The closest living relatives of the Elomeryx are bovids, suids, and cetaceans.
Anthracotheriidae is a paraphyletic family of extinct, hippopotamus-like artiodactyl ungulates related to hippopotamuses and whales. The oldest genus, Elomeryx, first appeared during the middle Eocene in Asia. They thrived in Africa and Eurasia, with a few species ultimately entering North America during the Oligocene. They died out in Europe and Africa during the Miocene, possibly due to a combination of climatic changes and competition with other artiodactyls, including pigs and true hippopotamuses. The youngest genus, Merycopotamus, died out in Asia during the late Pliocene, possibly for the same reasons. The family is named after the first genus discovered, Anthracotherium, which means "coal beast", as the first fossils of it were found in Paleogene-aged coal beds in France. Fossil remains of the anthracothere genus were discovered by the Harvard University and Geological Survey of Pakistan joint research project (Y-GSP) in the well-dated middle and late Miocene deposits of the Pothohar Plateau in northern Pakistan.
Dichobunidae is an extinct family of basal artiodactyl mammals from the early Eocene to late Oligocene of North America, Europe, and Asia. The Dichobunidae include some of the earliest known artiodactyls, such as Diacodexis.
Teilhardina was an early marmoset-like primate that lived in Europe, North America and Asia during the Early Eocene epoch, about 56-47 million years ago. The paleontologist George Gaylord Simpson is credited with naming it after the French paleontologist, Jesuit and philosopher Teilhard de Chardin.
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.
Duerotherium is an extinct genus of Paleogene 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.
The San Jose Formation is an Early Eocene geologic formation in the San Juan Basin of New Mexico and Colorado.
The Las Tetas de Cabra Formation is a geologic formation in Mexico. It preserves fossils dating back to the Wasatchian of the Early Eocene period.
The Lignites de Soissonais is a geologic formation in the Var, Marne departments of France. It preserves fossils dating back to the Ypresian stage of the Eocene period.
Homacodontidae is an extinct family of basal artiodactyl mammals from the early Eocene to late Oligocene of North America, Europe, and Asia.
Choeropotamidae, also known as Haplobunodontidae, are a family of extinct mammals, extinct herbivores, belonging to artiodactyls. They lived between the lower/middle Eocene and lower Oligocene and their remains were found in Europe and Africa.
The Silveirinha Formation is an Early Eocene geologic formation of the Mondego Basin in the Região Centro of central-western Portugal. The sandstones, siltstones and conglomerates were deposited in an alluvial environment.
The Alai Beds is an Early Eocene geologic formation in the Osh Region of southwestern Kyrgyzstan. The formation has provided many fossils of mammals, lizards, turtles and snakes.
Ancodonta is an infraorder of artiodactyl ungulates including modern hippopotamus and all mammals closer to hippos than to cetaceans (whales). Ancodonts first appeared in the Middle Eocene, with some of the earliest representatives found in fossil deposits in Southeast Asia. Throughout their evolutionary history they have occupied different browsing and grazing niches in North America, Eurasia and Africa. The last continent is notable as they were among the first laurasiatherian mammals to have migrated to Africa from Europe, where they competed with the native afrothere herbivores for the same niches. Of the nearly 50 genera that have existed, only two of them are extant – Choeropsis and Hippopotamus. The interrelationships within the ancodonts has been contended. The traditional notion is that there at minimum two families Anthracotheriidae and Hippopotamidae and were merely sister taxa. However many detailed research of the dentition among ancodonts, as well as how some anthracotheres were similar to hippos in appearance, lead the current consensus where Anthracotheriidae is paraphyletic to Hippopotamidae. Among the anthracotheres members of Bothriodontinae are among the closest to the ancestry of hippos, with the Oligocene aged Epirigenys from Lokon, Turkana, Kenya being the sister taxon to hippos. In response of this many similar clade names have been used for this clade.
Robiatherium is an extinct genus of Paleogene artiodactyls containing one species R. cournovense. The genus name derives from the locality of Robiac in France where some of its fossil were described plus the Greek θήρ/therium meaning "beast" or "wild animal." It was known only from the middle Eocene and, like other anoplotheriids, was endemic to western Europe. The genus was erected by Jean Sudre in 1988 for a species originally attributed to the xiphodont genus Paraxiphodon in 1978. Robiatherium had dentitions typical of the subfamily Anoplotheriinae, differing from other genera by specific differences in the molars. It is one of the earliest-appearing anoplotheriine species in the fossil record as well as the earliest to have appeared in central Europe.
