Metatheria Temporal range: | |
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Lycopsis longirostris , an extinct sparassodont, a relative of the marsupials | |
A mouse opossum ( Marmosa ) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Subclass: | Theria |
Clade: | Metatheria Huxley, 1880 |
Subgroups | |
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Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well as many extinct non-marsupial relatives. It is one of two groups placed in the clade Theria alongside Eutheria, which contains the placentals. Remains of metatherians have been found on all of Earth’s continents.
Distinctive characteristics (synapomorphies) of Metatheria include: a prehensile tail, the development of a capitular tail on the humerus, the loss of tooth replacement on the 2nd and 5th premolars and the retention of decidious teeth on the lower fifth premolars, the lower canines outwardly diverge from each other, the angular process on the dentary is equal to or less than half the length of the ramus, the dentary has a posterior masseteric shelf, and the lower 5th premolar has a "very trenchant" cristid obliqua/ectolophid. The permanent deciduous lower 5th premolars are molar like and were historically identified as 1st molars, with the third premolar found in basal therians being lost, leaving 4 premolars in the halves of each jaw. [4]
The relationships between the three extant divisions of mammals (monotremes, marsupials, and placentals) was long a matter of debate among taxonomists. [5] Most morphological evidence comparing traits, such as the number and arrangement of teeth and the structure of the reproductive and waste elimination systems, favors a closer evolutionary relationship between marsupials and placental mammals than either has with the monotremes, as does most genetic and molecular evidence. [6]
The earliest possible known metatherian is Sinodelphys szalayi , which lived in China during the Early Cretaceous around 125 million years ago (mya). [7] This makes it a contemporary to some early eutherian species that have been found in the same area. [8] However, Bi et al. (2018) reinterpreted Sinodelphys as an early member of Eutheria. The oldest uncontested metatherians are now 110 million year old fossils from western North America. [3] Metatherians were widespread in Asia and North America during the Late Cretaceous, including both Deltatheroida and Marsupialiformes, [9] with fossils also known from Europe during this time. During the Late Cretaceous, metatherians were more diverse than eutherians in North America. [4] Metatherians underwent a severe decline during the Cretaceous–Paleogene extinction event, more severe than that suffered by contemporary eutherians and multituberculates, and were slower to recover diversity. [9]
Morphological and species diversity of metatherians in Laurasia remained low in comparison to eutherians throughout the Cenozoic. [10] The two major groups of Cenozoic Laurasian metatherians, the opossum-like herpetotheriids and peradectids persisted into the Miocene before becoming extinct, with the North American herpetotheriid Herpetotherium , the European herpetotheriid Amphiperatherium and the peradectids Siamoperadectes and Sinoperadectes from Asia being the youngest Laurasian non-marsupial metatherians (with marsupials invading North America during the Pliocene-Pleistocene as part of the Great American interchange). [11] [9] Metatherians first arrived in Afro-Arabia during the Paleogene, probably from Europe, including the possible peradectoid Kasserinotherium from the Early Eocene of Tunisia and the herpetotheriid Peratherium africanum from the Early Oligocene of Egypt and Oman. The youngest African metatherian is the possible herpetotheriid Morotodon from the late Early Miocene of Uganda. [12] [13]
Metatherians arrived in South America from North America during the latest Cretaceous or Paleocene and underwent a major diversificiation, with South American metatherians including both the ancestors of extant marsupials as well as the extinct Sparassodonta, which were major predators in South American ecosystems during most of the Cenozoic, up until their extinction in the Pliocene, as well as the Polydolopimorphia, which likely had a wide range of diets. [10] Metatherians then declined in diversity in South America during the Late Eocene as well as the later Oligocene epoch. The oldest known Australian marsupials are from the early Eocene, and are thought to have arrived in the region after having dispersed via Antarctica from South America. [10] During the Oligocene epoch, Australian metatherians radiated rapidly, which contributed most to the global peak in metatherian diversity during the Early Miocene. The only known Antarctic metatherians are from the Early Eocene La Meseta Formation of the Antarctic Peninsula, where they are the most diverse group of mammals, and include marsupials as well as polydolopimorphians. [10]
Below is a metatherian cladogram from Wilson et al. (2016): [14]
Metatheria |
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Cladogram after [15] :
Metatheria | |
Below is a listing of metatherians that do not fall readily into well-defined groups.
Basal Metatheria
Ameridelphia incertae sedis:
Marsupialia incertae sedis:
Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of the defining features of marsupials is their unique reproductive strategy, where the young are born in a relatively undeveloped state and then nurtured within a pouch on their mother's abdomen.
Multituberculata is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late Cretaceous and Paleocene. They eventually declined from the mid-Paleocene onwards, disappearing from the known fossil record in the late Eocene. They are the most diverse order of Mesozoic mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as crown mammals outside either of the two main groups of living mammals—Theria, including placentals and marsupials, and Monotremata—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and Gondwanatheria as part of Allotheria.
Eomaia is a genus of extinct fossil mammals containing the single species Eomaia scansoria, discovered in rocks that were found in the Yixian Formation, Liaoning Province, China, and dated to the Barremian Age of the Lower Cretaceous about 125 million years ago. The single fossil specimen of this species is 10 centimetres (3.9 in) in length and virtually complete. An estimate of the body weight is 20–25 grams (0.71–0.88 oz). It is exceptionally well-preserved for a 125-million-year-old specimen. Although the fossil's skull is squashed flat, its teeth, tiny foot bones, cartilages and even its fur are visible.
Eutheria, also called Pan-Placentalia, is the clade consisting of placentals and all therian mammals that are more closely related to placentals than to marsupials.
Theria is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians and the metatherians but excludes the egg-laying monotremes and various extinct mammals evolving prior to the common ancestor of placentals and marsupials.
Sinodelphys is an extinct mammal from the Early Cretaceous, estimated to be 125 million years old. It was discovered and described in 2003 in rocks of the Yixian Formation in Liaoning Province, China, by a team of scientists including Zhe-Xi Luo and John Wible. While initially suggested to be the oldest known metatherian, later studies interpreted it as a eutherian.
Sparassodonta is an extinct order of carnivorous metatherian mammals native to South America, related to modern marsupials. They were once considered to be true marsupials, but are now thought to be a separate side branch that split before the last common ancestor of all modern marsupials.
The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked like mammals. The lineage leading to today's mammals split up in the Jurassic; synapsids from this period include Dryolestes, more closely related to extant placentals and marsupials than to monotremes, as well as Ambondro, more closely related to monotremes. Later on, the eutherian and metatherian lineages separated; the metatherians are the animals more closely related to the marsupials, while the eutherians are those more closely related to the placentals. Since Juramaia, the earliest known eutherian, lived 160 million years ago in the Jurassic, this divergence must have occurred in the same period.
Herpetotherium is an extinct genus of metatherian mammal, belonging to the possibly paraphyletic family Herpetotheriidae. Native to North America from the Eocene to Early Miocene, fossils have been found in California, Oregon, Texas, Florida, Montana, Wyoming, Colorado, North and South Dakota, Nebraska, and Saskatchewan. The oldest species, H. knighti, is dated to around 50.3 mya, and the most recent, an unnamed species, may be as recent as 15.97 mya. A morphological analysis of marsupials and basal metatherians conducted in 2007 found Herpetotherium to be the sister group to extant marsupials. It is the youngest known metatherian from North America until the migration of the Virginia opossum from South America within the last 2 million years.
Herpetotheriidae is an extinct family of metatherians, closely related to marsupials. Species of this family are generally reconstructed as terrestrial, and are considered morphologically similar to modern opossums. They are suggested to have been insectivores. Fossils of herpetotheriids come from North America, Asia, Europe, Africa, and perhaps South America. The oldest representative is Maastrichtidelphys from the latest Cretaceous (Maastrichtian) of the Netherlands and the youngest member is Amphiperatherium from the Middle Miocene of Europe. The group has been suggested to be paraphyletic, with an analysis of petrosal anatomy finding that North American Herpetotherium was more closely related to marsupials than the European Peratherium and Amphiperatherium.
Dryolestida is an extinct order of mammals, known from the Jurassic and Cretaceous. They are considered basal members of the clade Cladotheria, close to the ancestry of therian mammals. It is also believed that they developed a fully mammalian jaw and also had the three middle ear bones. Most members of the group, as with most Mesozoic mammals, are only known from fragmentary tooth and jaw remains.
This paleomammalogy list records new fossil mammal taxa that were described during the year 2011, as well as notes other significant paleomammalogy discoveries and events which occurred during that year.
Siamoperadectes is a genus of non-marsupial metatherian from the Miocene of Thailand. A member of Peradectidae, it is the first member of its clade known from South Asia, and among the last non-marsupial metatherians.
Groeberiidae is a family of strange non-placental mammals from the Eocene and Oligocene epochs of Patagonia, Argentina and Chile, South America. Originally classified as paucituberculate marsupials, they were suggested to be late representatives of the allothere clade Gondwanatheria. However, the relationship of the type genus, Groeberia, to Gondwanatheria has been firmly rejected by other scholars.
Anatoliadelphys maasae is an extinct genus of predatory metatherian mammal from the Eocene of Anatolia. It was an arboreal, cat-sized animal, with powerful crushing jaws similar to those of the modern Tasmanian devil. Although most mammalian predators of the northern hemisphere in this time period were placentals, Europe was an archipelago, and the island landmass now forming Turkey might have been devoid of competing mammalian predators, though this may not matter since other carnivorous metatherians are also known from the Cenozoic in the Northern Hemisphere. Nonetheless, it stands as a reminder that mammalian faunas in the Paleogene of the Northern Hemisphere were more complex than previously thought, and metatherians did not immediately lose their hold as major predators after their success in the Cretaceous.
The Itaboraí Formation is a highly fossiliferous geologic formation and Lagerstätte of the Itaboraí Basin in Rio de Janeiro, southeastern Brazil. The formation reaching a thickness of 100 metres (330 ft) is the defining unit for the Itaboraian South American land mammal age (SALMA), dating to the Early Eocene, approximately 53 to 50 Ma.
Gypsonictops is an extinct genus of leptictidan mammals of the family Gypsonictopidae, which was described in 1927 by George Gaylord Simpson. Species in this genus were small mammals and the first representatives of the order Leptictida, that appeared during the Upper Cretaceous.
Holoclemensia is an extinct genus of mammal of uncertain phylogenetic placement. It lived during the Early Cretaceous and its fossil remains were discovered in Texas.
Ambolestes is an extinct genus of eutherian mammal from the Early Cretaceous of China. It includes a single species, Ambolestes zhoui, known from a single complete skeleton recovered from the Yixian Formation, part of the fossiliferous Jehol biota. Ambolestes is one of the most basal eutherians, presenting a combination of features from both early eutherians (stem-placentals) and early metatherians (stem-marsupials). This is responsible for the generic name of Ambolestes: "ambo" is Latin for "both", while "-lestes" is a popular suffix for fossil mammals. The species name honors influential Jehol paleontologist Zhou Zhonghe.
Peradectidae is a family of small metatherian mammals, spanning from the Paleocene to the Miocene. Fossils are known from the Northern Hemisphere, including Europe, Asia and North America. The monophyly of the group has been questioned, with some authors suggesting that Peradectes should be the only genus placed in the family. The morphology of peradectids has been considered to be similar to opossums, with at least some exhibiting morphology suggesting a tree dwelling arboreal/scansorial lifestyle. Their diet is suggested to have included insects and fruit.