This article relies largely or entirely on a single source .(March 2023) |
Cokotherium Temporal range: Early Cretaceous, | |
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Holotype fossil of Cokotherium (a), with closer views of the skull (b), forelimbs (c), and CT-scanned 3D models of the skull (d-e) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Clade: | Eutheria |
Genus: | † Cokotherium Wang et al., 2022 |
Type species | |
†Cokotherium jiufotangensis Wang et al., 2022 |
Cokotherium is an extinct genus of eutherian mammal from the Early Cretaceous of China. It includes a single species, Cokotherium jiufotangensis, known from a single partial skeleton, missing a portion of the hindlimbs and tail. It was recovered from the Jiufotang Formation (120 Ma), the upper part of the fossiliferous Jehol biota. The generic name of Cokotherium honors the nickname of the late paleontologist Chuan-Kui Li, a specialist on the Jiufotang Formation. The specific name refers to the formation in question. Cokotherium is one of the youngest and most well-preserved Early Cretaceous eutherians, illustrating an array of transitional conditions between Early Cretaceous and Late Cretaceous members of Eutheria. [1]
The incisors, though conical and procumbent like other Early Cretaceous eutherians, are reduced in number; there are 4 in the upper jaw and 3 in the lower. Both the upper and lower canines are single-rooted, similar to metatherians (marsupial-line therians). Like most Mesozoic eutherians, Cokotherium has 5 premolars and 3 molars in both the upper and lower jaw. Similar to Late Cretaceous taxa, the last upper premolar is "molarized", with three roots and swollen cusps akin to the succeeding molars. Conversely, the protocone of the last upper premolar is much smaller than in the molars, and the last lower premolar lacks any molar-like specializations. The tribosphenic molars are simpler than in most Late Cretaceous eutherians. The upper molars have few additional protuberances or ridges beyond the three main cusps (protocone, paracone, and metacone). Likewise, the lower molars have a stout trigonid region (with three cusps: the protoconid > metaconid > paraconid) and an expansive talonid region (with two cusps: an entoconid and hypoconulid). [1]
The single known specimen of Cokotherium was in the process of replacing its third upper premolar, since both a deciduous tooth and a permanent tooth occupy the same position in the jaw. The other premolars are fully developed, including the second and fourth upper premolars. Alternating tooth replacement, with odd-numbered tooth replacement disjointed from their even-numbered neighbors, has been reported in a few other Mesozoic mammals, including the Jurassic eutherian Juramaia . It contrasts with the condition in placentals (living eutherians), which replace their teeth in a consistent sequence from front-to-back. The left lower canine also has yet to erupt, despite the full development and usage of the molars and remaining canines. The delayed eruption of the left lower canine is likely a consequence of localized injury rather than a developmental quirk. [1]
Cokotherium is the only eutherian with direct preservation of an ossified meckelian cartilage, similar to non-therian mammals. A few other early eutherians retain a meckelian sulcus, a groove on the ectotympanic which hinted that a meckelian cartilage was present at the base of Eutheria. The meckelian cartilage of Cokotherium is reduced to a slender strand lying against the angular process of the mandible, fully detached from the middle ear ossicles. The angular process itself is distinctly longer than in other early eutherians. Micro-CT scanning has revealed that Cokotherium has the most well-preserved osseous labyrinth (inner ear) in any Early Cretaceous eutherian, showing a combination of advanced and plesiomorphic ("primitive") features relative to later eutherians. [1]
Phylogenetic analyses by Wang et al. (2022) placed Cokotherium in a clade with Ambolestes and Sinodelphys at the base of Eutheria. In maximum parsimony analyses, Montanalestes also joins the clade, though it lies further down the eutherian stem in bayesian analyses. [1]
Phylogeny after Yang & Yang, 2023. [2]
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.
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.
Symmetrodonta is a group of Mesozoic mammals and mammal-like synapsids characterized by the triangular aspect of the molars when viewed from above, and the absence of a well-developed talonid. The traditional group of 'symmetrodonts' ranges in age from the latest Triassic to the Late Cretaceous, but most research in the last 20-30 years has concluded that they are not a true taxonomic group, but include several unrelated branches of the mammal tree. Despite this, the name is still used informally by some researchers for convenience, usually restricted to the spalacotheriids and zhangheotheriids.
Akidolestes is an extinct genus of mammals of the family Spalacotheriidae, a group of mammals related to therians.
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.
A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth. The concept is also used with regard to the leaflets of the four heart valves. The mitral valve, which has two cusps, is also known as the bicuspid valve, and the tricuspid valve has three cusps.
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.
Gobiconodon is an extinct genus of carnivorous mammals belonging to the family Gobiconodontidae. Undisputed records of Gobiconodon are restricted to the Early Cretaceous of Asia and North America, but isolated teeth attributed to the genus have also been described from formations in England and Morocco dating as far back as the Middle Jurassic. Species of Gobiconodon varied considerably in size, with G. ostromi, one of the larger species, being around the size of a modern Virginia opossum. Like other gobiconodontids, it possessed several speciations towards carnivory, such as shearing molariform teeth, large canine-like incisors and powerful jaw and forelimb musculature, indicating that it probably fed on vertebrate prey. Unusually among predatory mammals and other eutriconodonts, the lower canines were vestigial, with the first lower incisor pair having become massive and canine-like. Like the larger Repenomamus there might be some evidence of scavenging.
Asioryctitheria is an extinct order of early eutherians.
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.
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.
Triconodon is a genus of extinct mammal from the Early Cretaceous of England and France with two known species: T. mordax and T. averianovi. First described in 1859 by Richard Owen, it is the type genus for the order Triconodonta, a group of mammals characterised by their three-cusped (triconodont) molar teeth. Since then, this "simplistic" type of dentition has been understood to be either ancestral for mammals or else to have evolved multiple times, rendering "triconodonts" a paraphyletic or polyphyletic assemblage respectively, but several lineages of "triconodont" mammals do form a natural, monophyletic group, known as Eutriconodonta, of which Triconodon is indeed part of.
Spalacotheriidae is a family of extinct mammals belonging to the paraphyletic group 'Symmetrodonta'. They lasted from the Early Cretaceous to the Campanian in North America, Europe, Asia and North Africa.
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.
Azilestes is a genus of probable zhelestid eutherian mammal, a family consisting of small herbivores, that was discovered in the early Maastrichtian Grès de Labarre Formation of France. It is a monotypic genus, with only type species A. ragei being known. Only one specimen, the holotype described in 2021, is known. It consists of a partial dentary with teeth.
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.
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.
Chaoyangodens is an extinct genus of eutriconodont mammal from the Early Cretaceous of China. It includes a single species, Chaoyangodens lii, known from a single complete skeleton recovered from the Dawangzhangzi bed of the Yixian Formation, part of the fossiliferous Jehol biota. Chaoyangodens was a moderate-sized Mesozoic mammal. The generic name refers to Chaoyang Prefecture while the specific name honors the collector of the fossil, Hai-Jun Li. Chaoyangodens is intermediate in age between Liaoconodon and a diverse fauna of eutriconodonts from older beds of the Yixian Formation. Like Liaoconodon, it is not easily equated with other eutriconodonts, since it bears distinctive dental traits relative to recognized eutriconodont subgroups.
Microtherulum is an extinct genus of eutherian mammal known from the Early Cretaceous Jiufotang Formation of China. It is one of the most earliest and most primitive eutherians.