Arboroharamiya
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''Arboroharamiya'' 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
mammaliaform Mammaliaformes ("mammalian forms") is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined a ...
from the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
Tiaojishan Formation The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian-Oxfordian (stage), Oxfordian stages). It is known for its Lagerstätte, exceptionally preser ...
of
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an Autonomous regions of China, autonomous region of China. Its border includes two-thirds of the length of China's China–Mongolia border, border with the country of Mongolia. ...
, China. ''Arboroharamiya'' belongs to a group of mammaliaforms called Haramiyida. The genus contains three species: ''A. jenkinsi'' (the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
), ''A. allinhopsoni'' and ''A. fuscus''. Based on the melanosomes preserved in its fur, ''Arboroharamiya'' likely had a uniformly dark-brown coloration, similar to that of other early mammaliaforms including '' Vilevolodon'' and ''
Megaconus ''Megaconus'' is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, ''Megaconus mammaliaformis'' was first described in the journal ''Nature'' in 2013. ''Mega ...
''. When ''Arboroharamiya'' was included in a phylogenetic analysis of early mammals and mammaliaforms, Haramiyida was found to be a group within Mammalia, the true mammals. However, in a different analysis which included ''Megaconus'', Haramiyida was placed outside Mammalia as a more basal ("primitive") group of mammaliaforms. The classification of ''Arboroharamiya'' and other haramiyidans as true mammals fits with what most previous studies have found, but since ''Arboroharamiya'' and ''Megaconus'' were not included in the same phylogenetic analysis, their position within Mammaliaformes remains uncertain.


Description

''Arboroharamiya'' is the largest known haramiyidan, estimated to have weighed about . It has several features in common with living mammals, including a lower jaw formed by a single bone, the dentary, and hands and feet that each have four fingers with three bones each and one finger with two bones. ''Arboroharamiya'' is unlike any modern mammal in having a lower jaw that can move up, down, and backward, but not forward. It has a rodent-like dentition with enlarged
incisors 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 ...
and
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, '' ...
and no canines, which is also seen in
Multituberculata Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct Order (biology), order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the M ...
, but this feature probably evolved independently in ''Arboroharamiya''. ''Arboroharamiya'' had a long tail that might have been prehensile, and very long fingers. Based on the shape of its teeth, ''Arboroharamiya'' might have been a granivore or an omnivore. It was an arboreal animal, with the specimens of two species (''A. allinhopsoni'' and ''A. fuscus'') preserving the patagia, suggesting that it was a glider. Analysis on the fur impressions of ''A. fuscus'' revealed high concentration of copper associated with eumelanin, indicative of uniformly dark-brown coloration with no evidence of color patterns, which was also the case in other early mammaliaforms from China: '' Vilevolodon'', ''
Megaconus ''Megaconus'' is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, ''Megaconus mammaliaformis'' was first described in the journal ''Nature'' in 2013. ''Mega ...
'', indeterminate docodonts (SDUST-V0006 and SDUST-V0007) and indeterminate
eutheria Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
n (SDUST-V0008).


Classification

''Arboroharamiya'' belongs to a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
or evolutionary grouping called Mammaliaformes, which includes mammals and their closest extinct relatives from the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
and
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
periods. Within Mammaliaformes, ''Arboroharamiya'' falls within the clade Haramiyida. Haramiyidans have been known since the 1840s, but only from fossilized teeth and a single partial lower jaw. However, several features of the teeth have shown for many years that haramiyidans are among the most basal of mammaliaforms. Recent phylogenetic analyses, or analyses of evolutionary relationships, differ on whether or not haramiyidans are true mammals in the crown group Mammalia (the clade including the most recent common ancestor of living mammals, and all its descendants). Some analyses place Haramiyida outside crown group Mammalia, meaning that they diverged from other mammaliaforms before the most recent common ancestor of living mammals appeared. Other analyses place Haramiyida within Mammalia as part of an extinct group of Mesozoic mammals called
Allotheria Allotheria (meaning "other beasts", from the Ancient Greek language, Greek , '–other and , '–wild animal) is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower ...
, which also includes Multituberculata. When ''Arboroharamiya'' itself was incorporated into a phylogenetic analysis, Haramiyida fell within Allotheria as a group within Mammalia. Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
or evolutionary tree from the analysis:


References

{{Taxonbar, from=Q14611019 Fossil taxa described in 2013 Middle Jurassic synapsids of Asia Euharamiyida Prehistoric cynodont genera Paleontology in Inner Mongolia Taxa named by Xiaoting Zheng Taxa named by Shundong Bi Taxa named by Xiaoli Wang (paleontologist) Taxa named by Jin Meng