Euharamiyida Temporal range: | |
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Life restoration of Shenshou | |
Skull of Xianshou | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Order: | † Haramiyida |
Clade: | † Euharamiyida Bi et al. 2014 |
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, [1] or part of an earlier divergence within the synapsid line. [2] [3] It is disputed whether or not they are related to the Haramiyids from the Late Triassic, such as Haramiyavia . [4] 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. [1] [5]
The position of euharamyidans is contested. They are either considered crown group mammals as members of Allotheria, related to multituberculates, or they are considered to stem-group mammals within the Mammaliaformes. [1] [3] The position is often dependent on the relationships of euharamiyids to the Late Triassic haramiyids such as Haramiyavia and Thomasia. In some studies, the two groups are recovered as unrelated. [6] [4]
Huttenlocker et al. 2018 [7] | Mao et al. 2022 [8] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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The following taxonomy follows Mao et al. (2022) [8] unless otherwise cited.
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.
Lambdopsalis bulla is an extinct multituberculate mammal from the Late Paleocene of China and Mongolia. It is placed within the suborder Cimolodonta and is a member of the superfamily Taeniolabidoidea. Fossil remains have been found in the Late Paleocene Nomogen and Khashat Formations in Nao-mugen and Bayn Ulan of China and Mongolia.
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.
Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa and Antarctica during the Upper Cretaceous through the Paleogene. Until recently, they were known only from fragmentary remains. They are generally considered to be closely related to the multituberculates and likely the euharamiyidians, well known from the Northern Hemisphere, with which they form the clade Allotheria.
Allotheria is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower molariform teeth equipped with longitudinal rows of cusps and enlarged incisors. Typically, the canine teeth are also lost. Allotheria includes Multituberculata, Gondwanatheria, and probably Haramiyida, although some studies have recovered haramiyidans to be basal mammaliaforms unrelated to multituberculates. Allotherians are often placed as crown group mammals, more closely related to living marsupials and placentals than to monotremes or eutriconodonts.
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.
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 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, and Tikitherium.
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. Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia. Other studies have disputed 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.
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.
Arboroharamiya is an extinct genus of early mammal 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. 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. Recent interpretations of its specimen suggest that it possessed patagia and was a glider.
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.
Haramiyavia is a genus of synapsid in the clade Haramiyida that existed about 200 million years ago in the Rhaetian stage of the Triassic. Like other haramiyidans, it was likely a non-mammalian mammaliaform. It contains a single species, H. clemmenseni from the Fleming Fjord Formation of Greenland, and has been assigned to the monogeneric family Haramiyaviidae.
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.
Kermackodon is an extinct genus of allotherian mammal, known from the Middle Jurassic of England. It combines features of multituberculates with those of euharamyidans. The remains of type species, K. multicuspis were collected from Kirtlington Quarry in Oxford, England, by a team lead from UCL led by Professor Kenneth Kermack after whom the taxon is named, from sediments of the Forest Marble Formation, dating to the Bathonian stage of the Middle Jurassic. The genus and species were named by Percy M. Butler and Jerry Hooker in 2005. The remains comprise a left upper molar (M2), a lower last premolar, initially considered a left but later considered more likely to be right (p4), and an incomplete non-last upper premolar. A second species, K. oxfordensis, from Kirtlington and also sediments of the White Limestone Formation at Woodeaton Quarry was assigned to the genus in 2022, originally placed in the separate genus Eleutherodon. A 2020 study considered it to be more closely related to mutlituberculates than to euharamiyidans, while the 2022 study considered it to be a member of Euharamiyida.