Haramiyida | |
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Lower jaw and restoration of Haramiyavia , a basal haramiyidan from the Triassic of Greenland | |
Skull of Xianshou , a euharamiyidan from the Jurassic of China | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Order: | † Haramiyida Hahn, Sigogneau-Russell & Wouters, 1989 |
Subgroups | |
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Haramiyida is a possibly paraphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. [2] 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. [3] Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia. [4] [5] Other studies have disputed this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead. [6] 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. [7]
Order †Haramiyida [8] [9] Hahn, Sigogneau-Russell & Wouters, 1989 [Haramiyoidea Hahn, 1973 sensu McKenna & Bell, 1997]
The relationships of haramiydans to other mammals and mammaliaform cynodonts are controversial and have been subject to numerous conflicting phylogenetic analysis results. Major unresolved questions are whether or not haramiyidans are more closely related to marsupials and placental mammals (Theria) than they are to monotremes (and thus inside the crown group of Mammalia), or whether all living mammals (including therians and monotremes) are more closely related to each other than to haramiyidans (and thus placing Haramiyida outside crown Mammalia) and whether or not haramiyidans are closely related to multituberculates, an important of group of Mesozoic mammaliaforms typically regarded as crown group mammals, as part of the group Allotheria. While many studies recover Triassic haramiyidans and Jurassic euharamiyidans as closely related, some phylogenetic studies have recovered them as unrelated, find the Triassic haramiyidians as non-mammalian cynodonts, while recovering the Euharamiyida as crown-group mammals closely related to multituberculates. [7]
Cladogram from Luo et al 2017, showing a monophyletic Haramiyida outside of crown Mammalia unrelated to Multituberculata: [14]
Mammaliamorpha |
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Cladogram from Han et al. 2017, showing a paraphyletic Haramiyida within crown Mammalia as ancestral to Multituberculata: [15]
Mammaliamorpha |
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Cladogram from Hoffmann et al. 2020, showing a diphyletic Haramiyida. [7]
Mammaliamorpha |
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Simplified cladogram from Mao et al. 2024, showing a paraphyletic Haramiyida closely related to Multituberculata outside of crown Mammalia: [16]
Mammaliamorpha |
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Haramiyids seem to have generally been herbivorous or omnivorous, possibly the first mammalian herbivores; however, the sole haramiyid tested in a study involving Mesozoic mammal dietary habits, Haramiyavia , ranks among insectivorous species. [17] At least some species were very good climbers and were similar to modern day squirrels; [18] and several others have more recently been reassessed as possibly arboreal. General arboreal habits might explain their rarity in the fossil record. [19]
Several euharamiyidans, Maiopatagium , Xianshou , Vilevolodon and Arboroharamiya , took it one step further and developed the ability to glide, having extensive membranes similar to those of modern colugos. In many of these taxa, the coracoid bones (absent in modern therians but present in many other mammal groups, albeit highly reduced) are remarkably large and similar to those of birds and pterosaurs, presumably due to impact stresses at landing. [20] [21]
Mammalian tooth marks on dinosaur bones may belong to Sineleutherus , suggesting that some haramiyidans scavenged on dinosaur remains. [22]
The fossils of Late Triassic Haramiyids are primarily known from Europe and Greenland, [7] while the fossils of Euharamiyids are primarily known from the Middle to Late Jurassic of Asia. [23] Remains of eleutherodontids from Europe are only known from isolated teeth. [7]
The youngest haramiyid fossil genus has been considered to be possibly be Avashishta bacharamensis from the Maastrichtian of India, [24] however, this has not been robustly assessed by phylogenetics. [7] The youngest definitive euharamiyidan is Cryoharamiya from the Early Cretaceous Batylykh Formation of Yakutia, Russia. [23]
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.
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.
Cynodontia is a clade of eutheriodont therapsids that first appeared in the Late Permian, and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extinct ancestors and close relatives (Mammaliaformes), having evolved from advanced probainognathian cynodonts during the Late Triassic.
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 (Theria) than to monotremes or eutriconodonts, though some studies place the entirety of Allotheria outside of crown Mammalia.
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.
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.
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.
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.
Gobiconodontidae is a family of extinct mammals that ranged from the mid-Jurassic to the early Late Cretaceous, though most common during the Early Cretaceous. The Gobiconodontids form a diverse lineage of carnivorous non-therian mammals, and include some of the best preserved Mesozoic mammal specimens.
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.
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.
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.
Theriimorpha is a clade of mammals defined as including all mammals more closely related to therians than to monotremes. Eutriconodonta is usually considered among the most basal members of this group, with other members more closely related to therians like Allotherians placed in the subclade Theriiformes, though Eutriconodonta has also been recovered as less closely related to therians than monotremes are in some analyses, placing them outside the crown group of Mammalia. The unusual Late Jurassic digging mammal Fruitafossor has also been suggested to be a basal theriimorph in some studies. The position of Allotheria within Theriimorpha is controversial. While many studies recover Allotheria within crown Mammalia as more closely related to therians than to monotreme as a member of the clade Theriiformes, some studies recover the group as outside of crown Mammalia.
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.
Denise Sigogneau-Russell is a French palaeontologist who specialises in mammals from the Mesozoic, particularly from France and the UK. She is currently based at the Muséum National d'Histoire Naturelle.