Cokotherium
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''Cokotherium'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
eutherian Eutheria (from Greek , 'good, right' and , 'beast'; ), 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. Eutherians are distingu ...
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
of
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. 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 The Jiufotang Formation () is an Early Cretaceous geological formation in Chaoyang, Liaoning which has yielded fossils of feathered dinosaurs, primitive birds, pterosaurs, and other organisms (see Jehol Biota). It is a member of the Jehol group. T ...
(120 Ma), the upper part of the fossiliferous
Jehol biota The Jehol Biota includes all the living organisms – the ecosystem – of northeastern China between 133 and 120 million years ago. This is the Lower Cretaceous ecosystem which left fossils in the Yixian Formation and Jiufotang Formation. These d ...
. 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 The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
members of Eutheria.


Description


Dentition

The
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s, though conical and
procumbent This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary ...
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
metatheria 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 wel ...
ns (
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
-line therians). Like most
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
eutherians, ''Cokotherium'' has 5
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s and 3
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
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 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 ...
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 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 ...
). Likewise, the lower molars have a stout
trigonid 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 ...
region (with three cusps: the
protoconid 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 ...
>
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 i ...
>
paraconid 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 an expansive
talonid 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 ...
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 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 ...
). The single known specimen of ''Cokotherium'' was in the process of replacing its third upper premolar, since both a
deciduous tooth Deciduous teeth or primary teeth, also informally known as baby teeth, milk teeth, or temporary teeth,Fehrenbach, MJ and Popowics, T. (2026). ''Illustrated Dental Embryology, Histology, and Anatomy'', 6th edition, Elsevier, page 287–296. are ...
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 ''Juramaia'' is an extinct genus of a therian mammal, possibly a very basal eutherian mammal, known from the Late Jurassic ( Oxfordian stage) or Early Cretaceous deposits of western Liaoning, China. It is a small shrew-like mammal weighing aro ...
''. It contrasts with the condition in
placentals Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
(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 In humans, the cartilaginous bar of the mandibular arch is formed by what are known as Meckel's cartilages (right and left; also known as Meckelian cartilages), above which the incus and malleus are located. Meckel's cartilage arises from the first ...
, similar to non-therian mammals. A few other early eutherians retain a meckelian sulcus, a groove on the
ectotympanic The ectotympanic, or tympanicum, is a bony structure found in all mammals, located on the tympanic part of the temporal bone, which holds the tympanic membrane (eardrum) in place. In catarrhine primates (including humans), it takes a tube-shape. ...
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 __NOTOC__ The angle of the mandible (a.k.a. gonial angle, Masseteric Tuberosity, and Masseteric Insertion) is located at the posterior border at the junction of the lower border of the ramus of the mandible. The angle of the mandible, which may ...
of the mandible, fully detached from the
middle ear ossicles The ossicles (also called auditory ossicles) are three irregular bones in the middle ear of humans and other mammals, and are among the smallest bones in the human body. Although the term "ossicle" literally means "tiny bone" (from Latin ''ossicu ...
. 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 The bony labyrinth (also osseous labyrinth or otic capsule) is the rigid, bony outer wall of the inner ear in the temporal bone. It consists of three parts: the vestibule of the ear, vestibule, semicircular canals, and cochlea. These are cavities ...
(
inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
) in any Early Cretaceous eutherian, showing a combination of advanced and plesiomorphic ("primitive") features relative to later eutherians.


Classification

Phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
by Wang et al. (2022) placed ''Cokotherium'' in a clade with ''
Ambolestes ''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 (126 Ma), part of the fossil ...
'' and ''
Sinodelphys ''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-X ...
'' at the base of Eutheria. In
maximum parsimony In phylogenetics and computational phylogenetics, maximum parsimony is an optimality criterion under which the phylogenetic tree that minimizes the total number of character-state changes (or minimizes the cost of differentially weighted charact ...
analyses, ''
Montanalestes ''Montanalestes'' is an extinct mammal known from the Cretaceous in North America. Classification Known from only six fossil teeth, ''Montanalestes'' is an example of the less well-preserved early eutherian Eutheria (from Greek , 'good, ri ...
'' also joins the clade, though it lies further down the eutherian stem in
bayesian analyses Bayesian inference ( or ) is a method of statistical inference in which Bayes' theorem is used to calculate a probability of a hypothesis, given prior evidence, and update it as more information becomes available. Fundamentally, Bayesian inferen ...
. 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