Panperissodactyla Temporal range: | |
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Collage of living perissodactyls, clockwise from left: plains zebra (Equus quagga), Indian rhinoceros (Rhinoceros unicornis) and Brazilian tapir (Tapirus terrestris) | |
Macrauchenia patachonica (Litopterna) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Mirorder: | Euungulata |
Clade: | Panperissodactyla Welker et al, 2015 [1] |
Subgroups | |
[see classification] | |
Synonyms | |
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Panperissodactyla ("all perissodactyls", alternatively spelled Pan-Perissodactyla [2] ) is a clade of ungulates containing living order Perissodactyla (odd-toed ungulates) and all extinct ungulates more closely related to Perissodactyla than to Artiodactyla (even-toed ungulates). [1]
Groups thought to belong to this clade include Anthracobunia (including the families Anthracobunidae and Cambaytheriidae) known from the Paleogene of the Indian subcontinent, [3] [4] as well as the South American native ungulate groups Litopterna and Notoungulata, both of which went extinct approximately 12,000 years ago. [1] Other South American native ungulate groups also possibly belong to the clade, but their placement is uncertain. [5] The enigmatic aquatic Desmostylia have also been suggested to be related to perissodactyls in some studies, though others recover them as members of afrotherian clade Tethytheria. [6] The Northern Hemisphere "condylarth" group Phenacodontidae has been placed as closely related to perissodactyls in some studies, though others recover it as unrelated. [1]
Panperissodactyla was coined in 2015, to include all mammals more closely related to living perissodactyls than to any other living mammals, following a proteomic study which found Notoungulata and Litopterna to be closely related to Perissodactyla. [1] Panperissodactyla is preferred over the older Mesaxonia which was originally coined by Othniel Charles Marsh in 1884, as equivalent to the modern Perissodactyla. Richard Owen's original definition of Perissodactyla included the modern members in addition to the now known to be unrelated Hyracoidea. The name Mesaxonia was resurrected by Martin S. Fischer in 1989 as a replacement for Perissodactyla. [4] Perissodactylamorpha a clade coined to include Anthracobunia and Perissodactyla, is thought to be a subgroup of Panperissodactyla, as Anthracobunia is suggested to be more closely related to Perissodactyla than Notoungulata and Litopterna are. [4]
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.
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 62.5 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.
Wastebasket taxon is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by either their designated members' often superficial similarity to each other, or their lack of one or more distinct character states or by their not belonging to one or more other taxa. Wastebasket taxa are by definition either paraphyletic or polyphyletic, and are therefore not considered valid taxa under strict cladistic rules of taxonomy. The name of a wastebasket taxon may in some cases be retained as the designation of an evolutionary grade, however.
Toxodon is an extinct genus of large ungulate native to South America from the Pliocene to the end of the Late Pleistocene. 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, high intelligence, lack of grasping hands 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.
Chalicotheriidae is an extinct family of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene to the Early Pleistocene. They are often called chalicotheres, a term which is also applied to the broader grouping of Chalicotherioidea. They are noted for their unusual morphology compared to other ungulates, such as their clawed forelimbs. Members of the subfamily Chalicotheriinae developed elongate gorilla-like forelimbs that are thought to have been used to grasp vegetation. They are thought to have been browsers on foliage as well as possibly bark and fruit.
South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals that were indigenous to South America from the Paleocene until the end of the Late Pleistocene. They represented a dominant element of South America's Cenozoic terrestrial mammal fauna prior to the arrival of living unguate groups in South America during the Pliocene and Pleistocene as part of the Great American 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 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 Pliocene or Middle Pleistocene to the end of the Late 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 as a retracted nasal region, which in Macrauchenia manifests as the nasal opening being on the top of the skull between the eye sockets. This has historically been argued to correspond to the presence of a tapir-like proboscis, though recent authors suggest a moose-like prehensile lip or a saiga antelope-like nose to filter dust are more likely.
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.
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.
Pan-Euungulata is a clade of placental mammals from grandorder Ferungulata, consisting of the taxon Euungulata [sensu stricto] and all taxa (species) more closely related to it than to any other living species.