Ferungulata Temporal range: | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Clade: | Scrotifera |
Grandorder: | Ferungulata Simpson, 1945 [1] |
Subgroups | |
[see classification] | |
Synonyms | |
Ferungulata ("wild beasts and ungulates") is a grandorder of placental mammals that groups together mirorder Ferae and clade Pan-Euungulata. It has existed in two guises, a traditional one based on morphological analysis and a revised one taking into account more recent molecular analyses. The Fereungulata is a sister group to the order Chiroptera (bats) and together they make clade Scrotifera.
According to a 2022 study by Anne E. Kort, members of grandorder Ferungulata, in addition to their genetic similarities, share common synapomorphy in the lumbar vertebrae (the S-shaped postzygapophysis). In extant ferungulate mammals, this is known only within members of order Artiodactyla. However, this feature is found also in many extinct members of Ferungulata, including hyaenodonts, oxyaenids, mesonychids, arctocyonids and the stem-relatives of perissodactyls (like Cambaytherium ). [4] [5] This synapomorphy suggests that this may be basal to all Ferungulata and secondarily lost in modern members, like carnivorans and crown Perissodactyla. Previous study has shown that these S-shaped zygapophyses prevent torsion between vertebrae. As said in this study, it is possible this feature evolved in response to a need for stabilization in posterior spine as ribs became reduced. Also, in this and several other 2022 studies the extinct eutherians pantodonts, tillodonts and genus Deltatherium (family Deltatheriidae) are recognised as crown-group placental mammals, [4] [6] [7] [8] who also possess S-shaped postzygapophysis like other members of Ferungulata. [4]
The traditional Ferungulata was established by George Gaylord Simpson in 1945. [1] It grouped together the extant orders Carnivora, Perissodactyla and Artiodactyla with the Tubulidentata and the superorder Paenungulata, as well as a number of orders known only from fossils.
Simpson established the grouping on the basis of morphological criteria, but this traditional understanding of Ferungulata has been challenged by a more recent classification, relying upon genetic criteria. [3] [9] [10] These studies separated his ungulate orders into two distinct placental groups, within Afrotheria and Laurasiatheria, respectively. The 'true' ungulates (mirorder Euungulata), Perissodactyla and Artiodactyla, are included in the revised group, along with the Carnivora, and with the addition of pangolins (order Pholidota), but the Tubulidentata and paenungulates are excluded. Although Simpson placed whales (Cetacea) in a separate cohort, recent evidence linking them to Artiodactyla would mean that they belong here as well. To reflect this difference, the revised clade is usually referred to as Fereungulata.
The karyotypes of ferungulates are highly conserved: "the fereungulatan ancestor chromosomes (n = 23 + X) ... differ from those of the antecedent scrotiferan ancestor by five rearrangements that occurred over 1 My (citation excluded). For the fereungulatan ancestor, we discovered four ancestral syntenies..." [11]
Point coloration has been observed in a wide variety of ferungulates, including cats, foxes, horses, and dogs.
Traditional classification: [1] | Revised classification: [3] [9] [12] [13] |
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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.
Placental mammals are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguished from monotremes and marsupials in that the fetus is carried in the uterus of its mother to a relatively late stage of development. The name is something of a misnomer, considering that marsupials also nourish their fetuses via a placenta, though for a relatively briefer period, giving birth to less-developed young, which are then nurtured for a period inside the mother's pouch. Placentalia represents the only living group within Eutheria, which contains all mammals that are more closely related to placentals than they are to marsupials.
Eutheria, also called Pan-Placentalia, is the clade consisting of placental mammals and all therian mammals that are more closely related to placentals than to marsupials.
Afrotheria is a superorder of placental 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.
The Euarchonta are a proposed grandorder of mammals: the order Scandentia (treeshrews), and its sister Primatomorpha mirorder, containing the Dermoptera or colugos and the primates.
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.
Ferae is a mirorder of placental mammals in grandorder Ferungulata, that groups together clades Pan-Carnivora and Pholidotamorpha.
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.
Zooamata is a proposal for a clade of mammals uniting the Ferae with the Perissodactyla.
Panperissodactyla is a clade of ungulates containing living order Perissodactyla and all extinct ungulates more closely related to Perissodactyla than to Artiodactyla.
Protungulatum is an extinct genus of eutherian mammals within extinct family Protungulatidae, and is possibly one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Cretaceous to early Paleocene.
Pegasoferae is a proposed clade of mammals based on genomic research in molecular systematics by Nishihara, Hasegawa and Okada (2006).
Paratheria is an obsolete term for a taxonomic group including the xenarthran mammals and various groups thought to be related to them. It was proposed by Oldfield Thomas in 1887 to set apart the sloths, anteaters, armadillos, and pangolins, usually classified as placentals, from both marsupial and placental mammals, an arrangement that received little support from other workers. When teeth of the extinct gondwanathere mammals were first discovered in Argentina in the 1980s, they were thought to be related to xenarthrans, leading to renewed attention for the hypothesis that xenarthrans are not placentals. However, by the early 1990s, gondwanatheres were shown to be unrelated to xenarthrans, and xenarthrans are still considered to be placentals.
Scrotifera is a clade of placental mammals that groups together grandorder Ferungulata, Chiroptera (bats), other extinct members and their common ancestors. The clade Scrotifera is a sister group to the order Eulipotyphla based on evidence from molecular phylogenetics, and together they make superorder Laurasiatheria. The last common ancestor of Scrotifera is supposed to have diversified ca. 73.1 to 85.5 million years ago.
Altungulata or Pantomesaxonia is an invalid clade (mirorder) of ungulate mammals comprising the perissodactyls, hyracoids, and tethytheres.
Preptotheria is a superorder of placental mammals proposed by McKenna & Bell in their classification of mammals.
Ocepeia is an extinct genus of afrotherian mammal that lived in present-day Morocco during the middle Paleocene epoch, approximately 60 million years ago. First named and described in 2001, the type species is O. daouiensis from the Selandian stage of Morocco's Ouled Abdoun Basin. A second, larger species, O. grandis, is known from the Thanetian, a slightly younger stage in the same area. In life, the two species are estimated to have weighed about 3.5 kg (7.7 lb) and 10 kg (22 lb), respectively, and are believed to have been specialized leaf-eaters. The fossil skulls of Ocepeia are the oldest known afrotherian skulls, and the best-known of any Paleocene mammal in Africa.
Zalambdalestidae is a clade of Asian eutherians occurring during the Cretaceous. Once classified as Glires, features like epipubic bones and various cranial elements have identified these animals as outside of Placentalia, representing thus a specialised clade of non-placental eutherians without any living descendants, and potentially rather different from modern placentals in at least reproductive anatomy.
Pholidotamorpha is a clade of placental mammals from mirorder Ferae that includes the order Pholidota and extinct order Palaeanodonta.
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.