''Cokotherium'' is an extinct genus of eutherian

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from the

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. 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.

Description

Dentition

The incisors, though conical and

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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

premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...

s 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 A paracone is a 1960s atmospheric reentry or spaceflight mission abort concept using an inflatable ballistic cone.metacone). Likewise, the lower molars have a stout trigonid region (with three cusps: the protoconid >

metaconid Many different terms have been proposed for features of the tooth crown in mammals. The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...

> paraconid) and an expansive talonid region (with two cusps: an

entoconid Many different terms have been proposed for features of the tooth crown in mammals. The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...

and hypoconulid). 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 Permanent teeth or adult teeth are the second set of teeth formed in diphyodont mammals. In humans and old world simians, there are thirty-two permanent teeth, consisting of six maxillary and six mandibular molars, four maxillary and four mandibul ...

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.

Other features

''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

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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 (

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) in any Early Cretaceous eutherian, showing a combination of advanced and plesiomorphic ("primitive") features relative to later eutherians.

Classification

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

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. Phylogeny after Yang & Yang, 2023.

References

{{Taxonbar, from=Q117022397 Prehistoric eutherians Early Cretaceous mammals of Asia Cretaceous mammals Fossil taxa described in 2022 Jiufotang fauna Prehistoric mammal genera