Shuotherium Temporal range: Middle Jurassic-Late Jurassic | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Family: | † Shuotheriidae |
Genus: | † Shuotherium Chow & Rich, 1982 |
Type species | |
†Shuotherium dongi | |
Species | |
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Shuotherium is a fossil mammaliaform known from Middle-Late Jurassic of the Forest Marble Formation of England, [1] and the Shaximiao Formation of Sichuan, China. [2] [3]
The original holotype is composed of a partial dentary and seven teeth (two which are incomplete). [2] The holotypes for other species of this genus are solely represented by isolated molars. Shuotherium, along with Pseudotribos has been placed in the family Shuotheriidae as a sister taxon of the Australosphenida (see, Yinotheria), [4] making it a relative of modern monotremes. However, some studies place it and other shuothereres as closer to therian mammals, [5] [6] and another outside of Mammalia altogether. [7]
In the lower molars, the talonid is situated in front of the trigonid, such a unique dental form is distinct from the typical tribosphenic pattern. In this "pseudotribosphenic" trait, the mesial cingulid is expanded to form a pseudotalonid, and its distal talonid is underdeveloped. It shares with Australosphenida a thin, slender lower jaw but differs from the non-monotreme Ausktribosphenida by having more developed postdentary trough. Its dental formula has been reconstructed as: p4, m3 (four premolars, three molars). [4]
Kielan-Jaworowska, Cifelli & Luo 2002 had this to say regarding the fossil: [3]
In our view, the most compelling evidence as to the affinities of Shuotherium lies in the structure of the last premolar, which shares striking similarities to that of Australosphenida
Lower molar structure of Shuotherium and Australosphenida is obviously quite different, and for this reason we do not place Shuotherium within this Gondwanan clade. Based on the limited evidence available, however, we suggest that Shuotherium is a viable sister-taxon to Australosphenida.
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 the mammal groups. The third molar of humans is sometimes vestigial.
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.
Morganucodon is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, Morganucodon is well represented by abundant and well preserved material. Most of this comes from Glamorgan in Wales, but fossils have also been found in Yunnan Province in China and various parts of Europe and North America. Some closely related animals (Megazostrodon) are known from exquisite fossils from South Africa.
Allotheria is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower molariform teeth equipped with longitudinal rows of cusps and enlarged incisors. Typically, the canine teeth are also lost. Allotheria includes Multituberculata, Gondwanatheria, and probably Haramiyida, although some studies have recovered haramiyidans to be basal mammaliaforms unrelated to multituberculates. Allotherians are often placed as crown group mammals, more closely related to living marsupials and placentals (Theria) than to monotremes or eutriconodonts, though some studies place the entirety of Allotheria outside of crown Mammalia.
Tribosphenida is a group (infralegion) of mammals that includes the ancestor of Hypomylos, Aegialodontia and Theria. It belongs to the group Zatheria. The current definition of Tribosphenida is more or less synonymous with Boreosphenida.
Docodonta is an order of extinct Mesozoic mammaliaforms. They were among the most common mammaliaforms of their time, persisting from the Middle Jurassic to the Early Cretaceous across the continent of Laurasia. They are distinguished from other early mammaliaforms by their relatively complex molar teeth. Docodont teeth have been described as "pseudotribosphenic": a cusp on the inner half of the upper molar grinds into a basin on the front half of the lower molar, like a mortar-and-pestle. This is a case of convergent evolution with the tribosphenic teeth of therian mammals. There is much uncertainty for how docodont teeth developed from their simpler ancestors. Their closest relatives may have been certain Triassic "symmetrodonts", namely Woutersia, Delsatia.
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 its original definition and in many subsequent studies, but its relationship with the relationship with other members has been disputed by some scholars.
Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack et al. in 1973 as a replacement name for the paraphyletic Triconodonta.
Pseudotribos is an extinct genus of mammaliaform that lived in Northern China during the Middle Jurassic some 165 million years ago, possibly more closely related to monotremes than to theria, although other studies indicate that these shuotheres are closer to therians than to monotremes. The only known specimen was found in the Daohugou Bed in Inner Mongolia.
Trechnotheria is a group of mammals that includes the therians and some fossil mammals from the Mesozoic Era. It includes both the extinct symmetrodonts and the living Cladotheria.
Shuotheriidae is a small family of Jurassic mammaliaforms whose remains are found in China, Great Britain 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, with a 2024 study suggesting that shuotheriids were closely related to Docodonta outside of the Mammalia crown group.
Kuehneotherium is an early mammaliaform genus, previously considered a holothere, that lived during the Late Triassic-Early Jurassic Epochs and is characterized by reversed-triangle pattern of molar cusps. Although many fossils have been found, the fossils are limited to teeth, dental fragments, and mandible fragments. The genus includes Kuehneotherium praecursoris and all related species. It was first named and described by Doris M. Kermack, K. A. Kermack, and Frances Mussett in November 1967. The family Kuehneotheriidae and the genus Kuehneotherium were created to house the single species Kuehneotherium praecursoris. Modeling based upon a comparison of the Kuehneotherium jaw with other mammaliaforms indicates it was about the size of a modern-day shrew between 4 and 5.5 g at adulthood.
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.
Gobiconodontidae is a family of extinct mammals that ranged from the mid-Jurassic to the early Late Cretaceous, though most common during the Early Cretaceous. The Gobiconodontids form a diverse lineage of carnivorous non-therian mammals, and include some of the best preserved Mesozoic mammal specimens.
Paritatodon is an extinct mammaliaform which existed in Kyrgyzstan and England during the Jurassic period. It was originally the holotype specimen of Shuotherium kermacki, but Martin and Averianov (2010) argued that it resembled the genus Itatodon (Docodonta) and so renamed it Paritatodon.
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.
Ichthyoconodon is an extinct genus of eutriconodont mammal from the Lower Cretaceous of Morocco. It is notable for having been found in a unique marine location, and the shape of its teeth suggests an unusual, potentially fish-eating ecological niche. Analysis suggests it is part of a group of gliding mammals that includes Volaticotherium.
Vilevolodon is an extinct, monotypic genus of volant, arboreal euharamiyids from the Oxfordian age of the Late Jurassic of China. The type species is Vilevolodon diplomylos. The genus name Vilevolodon references its gliding capabilities, Vilevol, while don is a common suffix for mammalian taxon titles. The species name diplomylos refers to the dual mortar-and-pestle occlusion of upper and lower molars observed in the holotype; diplo, mylos.
Zhe-Xi Luo is an American paleontologist of Chinese origin, specializing in vertebrate paleontology, particularly mammal evolution, morphology, and systematics.