Tribosphenidans Temporal range: Late Jurassic to Holocene, | |
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Eomaia fossil | |
Kangaroo with her joey | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Clade: | Zatheria |
Clade: | Tribosphenida McKenna, 1975 |
Clades | |
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Synonyms | |
Boreosphenida Luo et al., 2001 |
Tribosphenida is a group (infralegion) of mammals that includes the ancestor of Hypomylos, Aegialodontia and Theria (the last common ancestor of marsupials and placentals plus all of its descendants). It belongs to the group Zatheria. [1] Current definition of Tribosphenida is more or less synonymous with Boreosphenida. [2] [3]
Tribosphenid mammals were originally grouped on the basis of triangular or V-shaped (tribosphenic) molars. Since then, other unrelated mammal groups have been found to have tribosphenic molars, such as the australosphenidans (a group that includes the still extant monotremes), suggesting that as a synapomorphy this is fundamentally useless as it evolved multiple times among mammals. [3]
However, a clade between the aforementioned groups, the "true Tribosphenida" or Boreosphenida, is still identifiable, united by characteristics such as the lack of a mesial cingulid and of a triangulated trigonid on the last premolar. [2] [3] They are also united by postcranial features such as the presence of a modern ear (though this too has evolved independently in many other groups, like monotremes [4] ), modern shoulder blades, and several features of the hindlimb. [2] [3]
Below is a cladogram showing one hypothesis of mammal relationships based on Rowe (1988) and McKenna and Bell (1997): [5] [6]
Mammaliaformes |
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Boreosphenida (from boreas, "northern wind" and sphen, "wedge") were early mammals that originated in the Northern Hemisphere and had tribosphenic molars (three-cusped cheek teeth). In boreosphenidans, the mandibular angle is placed posteriorly and the primitive postdentary trough (hole in the mandible) is absent (in contrast to Kuehneotheriidae, Eupantotheria, and Australosphenida.) They share the tribosphenic molars with the Australosphenida but differ from them by having cingulid cuspules but lacking a continuous mesial cingulid. Boreosphenidans also lack the triangulated trigonid on the last premolar found in Early Cretaceous mammals. They differ from Shuotherium (a monotreme-relative) in having the talonid placed posterior to the trigonid (like in modern tribosphenic mammals) in the lower molars, but upper molars similar to those of Shuotherium. [3]
The oldest boreosphenidans are from the Berriasian (~145-140 mya). They were restricted to the Northern Hemisphere during the Early Cretaceous, but spread to South America and India during the end of the Cretaceous. [3]
An alternative hypothesis is offered by Flannery and colleagues, who view australosphenidans as members of Tribosphenida (excluding monotremes, which they consider to be an unrelated group), with Barremian-Aptian Australian Bisphopidae as the sister taxon to all Northern Hemisphere Theria (Boreosphenida). [7] [8]
The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name molar derives from Latin, molaris dens, meaning "millstone tooth", from mola, millstone and dens, tooth. Molars show a great deal of diversity in size and shape across mammal groups. The third molar of humans is sometimes vestigial.
Cimolodonta is a clade of multituberculate mammals that lived from the Cretaceous to the Eocene. They probably lived something of a rodent-like existence until their ecological niche was assumed by true rodents. The more basal multituberculates are found in a different suborder, "Plagiaulacida", a paraphyletic group containing all non cimolodontan multituberculates.
Prototheria is an obsolete subclass of mammals which includes the living Monotremata and to which a variety of extinct groups, including Morganucodonta, Docodonta, Triconodonta and Multituberculata, have also been assigned. It is today no longer considered a valid grouping, but rather a paraphyletic evolutionary grade of basal mammals and mammaliaform cynodonts.
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.
Kollikodon is an extinct species of mammal, considered to be an early monotreme. It is known only from an opalised dentary fragment, with one premolar and two molars in situ, as well as a referred maxillary fragment containing the last premolar and all four molars. The fossils were found in the Griman Creek Formation at Lightning Ridge, New South Wales, Australia, as was Steropodon. Kollikodon lived in the Late Cretaceous period, during the Cenomanian age.
Steropodon is a genus of prehistoric platypus-like monotreme, or egg-laying mammal. It contains a single species, Steropodon galmani, that lived about 105 to 93.3 million years ago (mya) during the Cretaceous period, from early to middle Cenomanian. It is one of the oldest monotremes discovered, and is one of the oldest Australian mammal discoveries.
Teinolophos is a prehistoric species of monotreme, or egg-laying mammal, from the Teinolophidae. It is known from four specimens, each consisting of a partial lower jawbone collected from the Wonthaggi Formation at Flat Rocks, Victoria, Australia. It lived during the late Barremian age of the Lower Cretaceous.
The Australosphenida are a clade of mammals, containing mammals with tribosphenic molars, known from the Jurassic to Mid-Cretaceous of Gondwana. Although they have often been suggested to have acquired tribosphenic molars independently from those of Tribosphenida, this has been disputed. Fossils of australosphenidans have been found from the Jurassic of Madagascar and Argentina, and Cretaceous of Australia and Argentina. Monotremes have also been considered a part of this group in many studies, but this is also disputed.
Ausktribosphenidae is an extinct family of australosphenidan mammals from the Early Cretaceous of Australia and mid Cretaceous of South America.
Deltatheroida is an extinct group of basal metatherians that were distantly related to modern marsupials. The majority of known members of the group lived in the Cretaceous; one species, Gurbanodelta kara, is known from the late Paleocene (Gashatan) of China. Their fossils are restricted to Central Asia and North America. This order can be defined as all metatherians closer to Deltatheridium than to Marsupialia.
Amphitheriidae is a family of Mesozoic mammals restricted to the Middle Jurassic of Britain, with indeterminate members also possibly known from the equivalently aged Itat Formation in Siberia and the Anoual Formation of Morocco. They were members of Cladotheria, more derived than members of Dryolestida, and forming a close relationship with Peramuridae. Amphitheriidae is the only family of the order Amphitheriida.
Shuotherium is a fossil mammal known from Middle-Late Jurassic of the Forest Marble Formation of England, and the Shaximiao Formation of Sichuan, China.
Cladotheria is a clade of mammals. It contains modern therian mammals and several extinct groups, such as the "dryolestoids", amphitheriids and peramurids. The clade was named in 1975 by Malcolm McKenna. In 2002, it was defined as a node-based taxon containing "the common ancestor of dryolestids and living therians, plus all its descendants". A different, stem-based definition was given in 2013, in which Cladotheria contains all taxa that are closer to Mus musculus than to the "symmetrodont" Spalacotherium tricuspidens.
Shuotheriidae is a small family of Jurassic mammals whose remains are found in China, England and possibly Russia. They have been proposed to be close relatives of Australosphenida, together forming the clade Yinotheria. However, some studies suggest shuotheres are closer to therians than to monotremes, or that australosphenidans and therians are more closely related to each other than either are to shuotheres.
Monotremes are mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified by structural differences in their brains, jaws, digestive tract, reproductive tract, and other body parts, compared to the more common mammalian types. Although they are different from almost all mammals in that they lay eggs, like all mammals, the female monotremes nurse their young with milk.
Ambondro mahabo is a mammal from the Middle Jurassic (Bathonian) Isalo III Formation of Madagascar. The only described species of the genus Ambondro, it is known from a fragmentary lower jaw with three teeth, interpreted as the last premolar and the first two molars. The premolar consists of a central cusp with one or two smaller cusps and a cingulum (shelf) on the inner, or lingual, side of the tooth. The molars also have such a lingual cingulum. They consist of two groups of cusps: a trigonid of three cusps at the front and a talonid with a main cusp, a smaller cusp, and a crest at the back. Features of the talonid suggest that Ambondro had tribosphenic molars, the basic arrangement of molar features also present in marsupial and placental mammals. It is the oldest known mammal with putatively tribosphenic teeth; at the time of its discovery it antedated the second oldest example by about 25 million years.
Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.
Yinotheria is a proposed basal subclass clade of crown mammals uniting the Shuotheriidae, an extinct group of mammals from the Jurassic of Eurasia, with Australosphenida, a group of mammals known from the Jurassic to Cretaceous of Gondwana, which possibly include living monotremes. Today, there are only five surviving species of monotremes which live in Australia and New Guinea, consisting of the platypus and four species of echidna. Fossils of yinotheres have been found in Britain, China, Russia, Madagascar and Argentina. Contrary to other known crown mammals, they retained postdentary bones as shown by the presence of a postdentary trough. The extant members (monotremes) developed the mammalian middle ear independently.
Teinolophidae is an extinct family of small, carnivorous mammals that were the earliest known monotremes and were endemic to what would become Australia. Two genera are known: Teinolophos, and possibly also Stirtodon.
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.
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