Perissodactyla is an order of ungulates. The order includes about 17 living species divided into three families: Equidae, Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs). They typically have reduced the weight-bearing toes to three or one of the five original toes, though tapirs retain four toes on their front feet. The nonweight-bearing toes are either present, absent, vestigial, or positioned posteriorly. By contrast, artiodactyls bear most of their weight equally on four or two of the five toes: their third and fourth toes. Another difference between the two is that perissodactyls digest plant cellulose in their intestines, rather than in one or more stomach chambers as artiodactyls, with the exception of Suina, do.
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.
Afrotheria is a superorder of mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephant shrews, otter shrews, tenrecs, aardvarks, hyraxes, elephants, sea cows, and several extinct clades. Most groups of afrotheres share little or no superficial resemblance, and their similarities have only become known in recent times because of genetics and molecular studies. Many afrothere groups are found mostly or exclusively in Africa, reflecting the fact that Africa was an island continent from the Cretaceous until the early Miocene around 20 million years ago, when Afro-Arabia collided with Eurasia.
Paenungulata is a clade of "sub-ungulates", which groups three extant mammal orders: Proboscidea, Sirenia, and Hyracoidea (hyraxes). At least two more possible orders are known only as fossils, namely Embrithopoda and Desmostylia.
Litopterna is an extinct order of South American native ungulates that lived from the Paleocene to the end of the Pleistocene-early Holocene around 63 million-12,000 years ago, and were also present in Antarctica during the Eocene. They represent the second most diverse group of South American ungulates after Notoungulata. It is divided into nine families, with Proterotheriidae and Macraucheniidae being the most diverse and last surviving families.
Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wastebasket taxon, having served as a dumping ground for classifying ungulates which had not been clearly established as part of either Perissodactyla or Artiodactyla, being composed thus of several unrelated lineages.
Toxodon is an extinct genus of large ungulate native to South America from the Late Miocene to early Holocene epochs. Toxodon is a member of Notoungulata, an order of extinct South American native ungulates distinct from the two living ungulate orders that had been indigenous to the continent for over 60 million years since the early Cenozoic, prior to the arrival of living ungulates into South America around 2.5 million years ago during the Great American Interchange. Toxodon is a member of the family Toxodontidae, which includes medium to large sized herbivores. Toxodon was one of the largest members of Toxodontidae and Notoungulata, with Toxodon platensis having an estimated body mass of 1,000–1,200 kilograms (2,200–2,600 lb).
Notoungulata is an extinct order of ungulates that inhabited South America from the early Paleocene to the end of the Pleistocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms resembling animals as disparate as rabbits and rhinoceroses. Notoungulata are the largest group of South American native ungulates, with over 150 genera in 14 families having been described, divided into two major subgroupings, Typotheria and Toxodontia. Notoungulates first diversified during the Eocene. Their diversity declined from the late Neogene onwards, with only the large toxodontids persisting until the end of the Pleistocene, perishing as part of the Late Pleistocene megafauna extinctions along with most other large mammals across the Americas. Collagen sequence analysis suggests that notoungulates are closely related to litopterns, another group of South American ungulates, and their closest living relatives being perissodactyls, including rhinoceroses, tapirs and equines as part of the clade Panperissodactyla. However their relationships to other South American ungulates are uncertain. Several groups of notoungulates separately evolved ever-growing cheek teeth.
Laurasiatheria is a superorder of placental mammals that groups together true insectivores (eulipotyphlans), bats (chiropterans), carnivorans, pangolins (pholidotes), even-toed ungulates (artiodactyls), odd-toed ungulates (perissodactyls), and all their extinct relatives. From systematics and phylogenetic perspectives, it is subdivided into order Eulipotyphla and clade Scrotifera. It is a sister group to Euarchontoglires with which it forms the magnorder Boreoeutheria. Laurasiatheria was discovered on the basis of the similar gene sequences shared by the mammals belonging to it; no anatomical features have yet been found that unite the group, although a few have been suggested such as a small coracoid process, a simplified hindgut and allantoic vessels that are large to moderate in size. The Laurasiatheria clade is based on DNA sequence analyses and retrotransposon presence/absence data. The superorder originated on the northern supercontinent of Laurasia, after it split from Gondwana when Pangaea broke up. Its last common ancestor is supposed to have lived between ca. 76 to 90 million years ago.
South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals of controversial affinities that were indigenous to South America prior to the Great American Biotic Interchange. They comprise five major groups conventionally ranked as orders—Astrapotheria, Litopterna, Notoungulata, Pyrotheria, and Xenungulata—as well as the primitive "condylarth" groups Didolodontidae and Kollpaniinae. It has been proposed that some or all of the members of this group form a clade, named Meridiungulata, though the relationships of South American ungulates remain largely unresolved. The two largest groups of South American ungulates, the notoungulates and the litopterns, were the only groups to persist beyond the mid Miocene. Only a few of the largest species of notoungulates and litopterns survived until the end-Pleistocene extinction event around 12,000 years ago where they became extinct with most other large mammals in the Americas, shortly after the first arrival of humans into the region.
Macrauchenia is an extinct genus of large ungulate native to South America from the late Pliocene to the end of the Pleistocene. It is a member of the extinct order Litopterna, a group of South American native ungulates distinct from the two orders which contain all living ungulates which had been present in South America since the early Cenozoic, over 60 million years ago, prior to the arrival of living ungulates in South America around 2.5 million years ago as part of the Great American Interchange. The bodyform of Macrauchenia has been described as similar to a camel, being one of the largest known litopterns, with an estimated body mass of around 1 tonne. The genus gives its name to its family, Macraucheniidae, which like Macrauchenia typically had long necks and three toed feet, as well a retracted nasal region, which in Macrauchenia manifests as the nasal opening being on the top of the skull behind the eye sockets. This has historically been argued to correspond to the presence of a tapir-like proboscis, but some recent authors suggest a moose-like prehensile lip or a saiga antelope-like nose to filter dust are more likely.
Macraucheniidae is a family in the extinct South American ungulate order Litopterna, that resembled various camelids. The reduced nasal bones of their skulls was originally suggested to have housed a small proboscis, similar to that of the saiga antelope. However, one study suggested that they were openings for large moose-like nostrils. Conversely, prehistoric pictographs by indigenous people seems to depict animals interpreted as macraucheniids with trunks. Their hooves were similar to those of rhinoceroses today, with a simple ankle joint and three digits on each foot. Thus, they may have been capable of rapid directional change when running away from predators, such as large phorusrhacid terror birds, sparassodont metatherians, giant short-faced bears (Arctotherium) and saber-toothed cats (Smilodon). Macraucheniids probably lived in large herds to gain protection against these predators, as well as to facilitate finding mates for reproduction.
Anthracobunidae is an extinct family of stem perissodactyls that lived in the early to middle Eocene period. They were originally considered to be a paraphyletic family of primitive proboscideans possibly ancestral to the Moeritheriidae and the desmostylians. The family has also thought to be ancestral to the Sirenia.
Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, 59 to 11.8 million years ago. Astrapotheres were large, rhinoceros-like animals and have been called one of the most bizarre orders of mammals with an enigmatic evolutionary history.
Toxodontidae is an extinct family of notoungulate mammals, known from the Oligocene to the Holocene of South America, with one genus, Mixotoxodon, also known from the Pleistocene of Central America and southern North America. Member of the family were medium to large-sized, ranging from around 350–400 kilograms (770–880 lb) in Nesodon to 1,000–1,200 kilograms (2,200–2,600 lb) in Toxodon, and had medium to high-crowned dentition, which in derived members of the group evolved into ever-growing cheek teeth. Isotopic analyses have led to the conclusion that Pleistocene members of the family were flexible mixed feeders.
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.
Altungulata or Pantomesaxonia is an invalid clade (mirorder) of ungulate mammals comprising the perissodactyls, hyracoids, and tethytheres.
Cambaytherium is an extinct genus of placental mammals in the family Cambaytheriidae whose fossils were found in an open pit coal mine located in Gujarat, India. The mine was a treasure trove full of teeth and bones, over 200 of which were identified as belonging to Cambaytherium thewissi. The fossils were dated to the Early Eocene, 54.5 million years ago, making them slightly younger than the oldest known fossils belonging to the order Perissodactyla.
Cambaytheriidae is a family of primitive four or five-toed ungulates native to the Indian subcontinent. They lived during the Early Eocene epoch and are distinguished by the presence of bunodont teeth suitable for eating tough vegetation. They are related to, but distinct from, the early perissodactyls, and may also be closely related to the anthracobunids as a sister group to the Perissodactyla.
Sparnotheriodontidae is an enigmatic extinct family of litopterns known primarily from teeth. Sparnotheriodontids are one of two South American native ungulate clades known to have reached Antarctica, the other being astrapotheres.
