Arboroharamiya Temporal range: Middle Jurassic, | |
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Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Order: | † Haramiyida |
Genus: | † Arboroharamiya Zheng et al., 2013 |
Species | |
†Arboroharamiya jenkinsi Contents |
Arboroharamiya is an extinct genus of early mammal (or possibly a non-mammalian mammaliaform) from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. Arboroharamiya belongs to a group of mammaliaforms called Haramiyida. The type species Arboroharamiya jenkinsi was described in the journal Nature in 2013 alongside a description of the closely related haramiyidan Megaconus . Unlike Megaconus, which is thought to have been ground-dwelling, Arboroharamiya was arboreal. [2] It has a long tail that might have been prehensile, and very long fingers. Based on the shape of its teeth, Arboroharamiya might have been an omnivore or a seed eater. [3] Recent interpretations of its specimen suggest that it possessed patagia and was a glider. [4]
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, Megaconus was included in a different phylogenetic analysis, and that analysis placed Haramiyida 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. [3]
Arboroharamiya is the largest known haramiyidan, estimated to have weighed about 354 g. 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 and molars and no canines. A rodent-like dentition is also seen in Multituberculata, an early group of mammals that might be closely related to Arboroharamiya, but it probably evolved independently in Arboroharamiya. [2]
Arboroharamiya belongs to a clade or evolutionary grouping called Mammaliaformes, which includes mammals and their closest extinct relatives from the Triassic and Jurassic 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, 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 or evolutionary tree from the analysis:
Mammaliaformes |
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Multituberculata is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late Cretaceous and Paleocene. They eventually declined from the mid Paleocene onwards, disappearing from the known fossil record in the late Eocene. They are the most diverse order of Mesozoic mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as crown mammals outside either of the two main groups of living mammals—Theria, including placentals and marsupials, and Monotremata—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and Gondwanatheria as part of Allotheria.
Synapsids are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptiles and birds. The group includes mammals and every animal more closely related to mammals than to sauropsids. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.
Hahnodon is an extinct genus of mammaliaforms from the Early Cretaceous Ksar Metlili Formation in Morocco. Although originally considered to be a relatively early member of the extinct clade Multituberculata, recent studies indicate that it instead is a haramiyid.
Hahnodontidae is a family of extinct mammaliaforms from Early Cretaceous deposits in Morocco and the Western United States. Although originally considered to belong to the extinct clade Multituberculata, recent work indicates that hahnodontids belong to the more primitive clade Haramiyida.
Castorocauda is an extinct, semi-aquatic, superficially otter-like genus of docodont mammaliaforms with one species, C. lutrasimilis. It is part of the Yanliao Biota, found in the Daohugou Beds of Inner Mongolia, China dating to the Middle to Late Jurassic. It was part of an explosive Middle Jurassic radiation of Mammaliaformes moving into diverse habitats and niches. Its discovery in 2006, along with the discovery of other unusual mammaliaforms, disproves the previous hypothesis of Mammaliaformes remaining evolutionarily stagnant until the extinction of the non-avian dinosaurs.
Docodonta is an order of extinct mammaliaforms that lived during the Mesozoic, from the Middle Jurassic to Early Cretaceous. They are distinguished from other early mammaliaforms by their relatively complex molar teeth, from which the order gets its name. Until recently, Docodonta were represented primarily by teeth and jaws found across former Laurasia,. However, recent discoveries in China include some exceptionally well preserved, almost complete body fossils.
Mammaliaformes is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent common ancestor of Morganucodonta and the crown group mammals; the latter is the clade originating with the most recent common ancestor of extant Monotremata, Marsupialia, and Placentalia. Besides Morganucodonta and the crown group mammals, Mammaliaformes includes Docodonta and Hadrocodium as well as the Triassic Tikitherium, the earliest known member of the group.
Tritylodontidae is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, bearing several mammalian traits like erect limbs, endothermy and details of the skeleton. They were the last-known family of the non-mammaliaform synapsids, persisting into the Early Cretaceous.
Volaticotherium antiquum is an extinct, gliding, insectivorous mammal that lived in Asia during the Jurassic period, around 164 mya. It is the only member of the genus Volaticotherium.
Volaticotherini is a clade of eutriconodont mammals from the Mesozoic. In addition to the type genus Volaticotherium, it includes the genera Argentoconodon, Ichthyoconodon, and potentially Triconolestes.
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.
Haramiyida is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remains, resemble those of the multituberculates. However, based on Haramiyavia, the jaw is less derived; and at the level of evolution of earlier basal mammals like Morganucodon and Kuehneotherium, with a groove for ear ossicles on the dentary. If they are early multituberculates, they would be the longest lived mammalian clade of all time. However, a more recent study in November 2015 may dispute this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead. It is also disputed whether the Late Triassic species are closely related to the Jurassic and Cretaceous members belonging to Euharamiyida/Eleutherodontida, as some phylogenetic studies recover the two groups as unrelated, recovering the Triassic haramiyidians as non-mammalian cynodonts, while recovering the Euharamiyida as crown-group mammals closely related to multituberculates.
Morganucodonta is an extinct order of basal Mammaliaformes, a group including crown-group mammals (Mammalia) and their close relatives. Their remains have been found in Southern Africa, Western Europe, North America, India and China. The morganucodontans were probably insectivorous and nocturnal, though like eutriconodonts some species attained large sizes and were carnivorous. Nocturnality is believed to have evolved in the earliest mammals in the Triassic as a specialisation that allowed them to exploit a safer, night-time niche, while most larger predators were likely to have been active during the day.
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. Megaconus is thought to have been a herbivore that lived on the ground, having a similar posture to modern-day armadillos and rock hyraxes. Megaconus was in its initial description found to be member of a group called Haramiyida. A phylogenetic analysis published along its description suggested that haramiyidans originated before the appearance of true mammals, but in contrast, the later description of the haramiyidan Arboroharamiya in the same issue of Nature indicated that haramyidans were true mammals. If haramiyidans are not mammals, Megaconus would be one of the most basal ("primitive") mammaliaforms to possess fur, and an indicator that fur evolved in the ancestors of mammals and not the mammals themselves. However, later studies cast doubt on the euharamiyidan intrepretation, instead finding it to be a basal allotherian mammal.
Shenshou is a genus of haramiyidan dating from the Oxfordian stage of the Late Jurassic, approximately 160 million years ago. Fossils were recovered from the Tiaojishan Formation in the Liaoning province of China.
Xianshou is a genus of gliding haramiyidan synapsid known from the Oxfordian stage of the Jurassic period, approximately 160 million years ago. Two species, X. linglong and X. songae, are known from fossils of the Tiaojishan Formation in the Liaoning province of China.
Euharamiyida also known as Eleutherodontida, is clade of early mammals or mammal-like cynodonts from the Middle Jurassic to Early Cretaceous of Eurasia and possibly North America. The group is sometimes considered a sister group to Multituberculata, or part of an earlier divergence within the synapsid line. It is disputed whether or not they are related to the Haramiyids from the Late Triassic, such as Haramiyavia. The morphology of their teeth indicates that they were herbivorous or omnivorous. Some members of the group are known to be arboreal, including gliding forms similar to modern flying squirrels or colugos.
Yi is a genus of scansoriopterygid dinosaurs from the Late Jurassic of China. Its only species, Yi qi, is known from a single fossil specimen of an adult individual found in Middle or Late Jurassic Tiaojishan Formation of Hebei, China, approximately 159 million years ago. It was a small, possibly tree-dwelling (arboreal) animal. Like other scansoriopterygids, Yi possessed an unusual, elongated third finger, that appears to have helped to support a membranous gliding plane made of skin. The planes of Yi qi were also supported by a long, bony strut attached to the wrist. This modified wrist bone and membrane-based plane is unique among all known dinosaurs, and might have resulted in wings similar in appearance to those of bats.
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.
Cifelliodon is an extinct genus of mammaliaforms from the Lower Cretaceous of North America. In the describing paper, it was considered one of the latest surviving haramiyids yet known, belonging to the family Hahnodontidae. Its discovery led to the proposal to remove hahnodontids from the larger well-known group, the multituberculates. However, later papers have considered it to be a basal allotherian outside of Haramiyida.