Paenungulatomorpha Temporal range: Paleocene - Recent, | |
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Crowned paenungulates and Ocepeia | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Superorder: | Afrotheria |
Clade: | Paenungulatomorpha Gheerbrant, 2016 |
Subgroups | |
Paenungulatomorpha is a clade of afrotherian mammals that can be characterized according to Gheerbrant et al. (2016) [1] :
by a mandibular retromolar fossa, the absence of hypocone, an ectoloph selenodont and linked to strong styles such as mesostyle in basal taxa, and a more or less developed pseudohypocone.
Originally it was thought paenungulates were closely related to the perissodactyls in the clade Altungulata due to shared attributes in the dental, osteological and soft-tissues. [2] [3] [4] However this view was contested as molecular studies have widely supported that paenungulates are not related to the perissodactyls, instead more related to the widely different clade Afroinsectiphilia, with them forming the clade Afrotheria. [5] [6] [7] [8] [9] There has been also some morphological data to support Afrotheria, [10] [11] [12] though the paleontological record to support was lacking until reexamination of the genera Ocepeia [13] and Abdounodus , [1] as well as the description of the basal Hadrogeneios in 2023 [14] has shown not only fossil evidence for the inclusion of paenungulates as afrotherians but also any morphological similarities between paenungulates and perissodactyls is due to convergence. [1]
Below is a cladogram from Gheerbrant (2023): [14]
Paenungulatomorpha | |
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.
The Desmostylia are an extinct order of aquatic mammals native to the North Pacific from the early Oligocene (Rupelian) to the late Miocene (Tortonian). Desmostylians are the only known extinct order of marine mammals.
Elephant shrews, also called jumping shrews or sengis, are small insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order Macroscelidea. Their traditional common English name "elephant shrew" comes from a perceived resemblance between their long noses and the trunk of an elephant, and their superficial similarity with shrews in the order Eulipotyphla. However, phylogenetic analysis has revealed that elephant shrews are not properly classified with true shrews, but are in fact more closely related to elephants than to shrews. In 1997, the biologist Jonathan Kingdon proposed that they instead be called "sengis", a term derived from the Bantu languages of Africa, and in 1998, they were classified into the new clade Afrotheria.
A tenrec is a mammal belonging to any species within the afrotherian family Tenrecidae, which is endemic to Madagascar. Tenrecs are a very diverse group; as a result of convergent evolution, some resemble hedgehogs, shrews, opossums, rats, and mice. They occupy aquatic, arboreal, terrestrial, and fossorial environments. Some of these species, including the greater hedgehog tenrec, can be found in the Madagascar dry deciduous forests. However, the speciation rate in this group has been higher in humid forests.
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.
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.
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.
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.
Embrithopoda ("heavy-footed") is an order of extinct mammals known from Asia, Africa and Eastern Europe. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late Paleocene to the late Eocene; however, the order is best known from its terminal member, the elephantine Arsinoitherium.
South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals of controversial affinities 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 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.
The Afroinsectiphilia is a clade that has been proposed based on the results of recent molecular phylogenetic studies. Many of the taxa within it were once regarded as part of the order Insectivora, but Insectivora is now considered to be polyphyletic and obsolete. This proposed classification is based on molecular studies only, and there is no morphological evidence for it.
Ferungulata 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.
Pseudoungulata, or "false hoofed mammals", is a possible clade made up of two subgroups, aardvarks and paenungulates. Before this group was proposed, it was thought that aardvarks were more closely related to xenarthrans. However, all of these mammals are now considered to be part of Afrotheria, which also includes elephant shrews and afrosoricidans. Other positions of aardvarks within Afrotheria are possible, such as being closest relatives of elephant shrews and/or afrosoricidans.
Ptolemaiida is a taxon of wolf-sized afrothere mammals that lived in northern and eastern Africa during the Paleogene. The oldest fossils are from the latest Eocene strata of the Jebel Qatrani Formation, near the Fayum oasis in Egypt. A tooth is known from an Oligocene-aged stratum in Angola, and Miocene specimens are known from Kenya and Uganda.
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.
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.
Abdounodus is an extinct genus of mammal known from the middle Paleocene of Northern Africa. The sole species, A. hamdii, is known from teeth and jaw bones discovered in the Ouled Abdoun Basin of present-day Morocco in 2001.
Hadrogeneios is an extinct genus of afrotherian mammal recovered from the Khouribga Phosphates of the Ouled Abdoun Basin dating from the Selandian to the Thanetian. Only the type species H. phosphaticus is known for this genus, with the material being of dental and jaw specimens. The placement of Hadrogeneios recovered it to be the most basal stem-paenungulate, the clade that includes elephants, sea cows, and hyraxes. Despite this Hadrogeneios was a contemporary of more derived members of the group such as Abdounodus, Ocepeia, and the early proboscideans Eritherium, Phosphatherium, and Daouitherium.
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