Morganucodonta Temporal range: | |
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Life restoration of a Megazostrodon , Natural History Museum, London | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Order: | † Morganucodonta Kermack, Mussett, & Rigney, 1973 |
Subgroups | |
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Morganucodonta ("Glamorgan teeth") 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. [1] Nocturnality is believed to have evolved in the earliest mammals in the Triassic (called the nocturnal bottleneck) 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 (though some dinosaurs, for example, were nocturnal as well [2] ).
Morganucodontans had a double jaw articulation made up of the dentary-squamosal joint as well as a quadrate-articular one. This implies that they also retained one of their postdentary bones: the articular. There is a trough at the back of the jaw on the inside (lingual) where the postdentary bones sat. The double jaw articulation and retention of the postdentary bones are characteristic of many of the earliest Mammaliaformes, but are absent today in mammals: All crown-group mammals have a lower jaw that is composed of a single bone, and the articular has been incorporated into the middle ear, having become the malleus (while the quadrate has become the incus).
Unlike more basal therapsids such as Sinoconodon , morganucodontan teeth were diphyodont (meaning that they replaced their teeth once, having a 'milk' set and 'adult' set as seen in today's mammals including humans [3] and not polyphyodont (meaning that the teeth are constantly replaced, and the animal and its teeth get larger throughout its lifetime, as in reptiles). Evidence of lactation is present in Wareolestes , via tooth replacement patterns. [4]
Morganucodontans also had specialised teeth – incisors, canines, molars and premolars – for food processing, rather than having similarly shaped teeth along the tooth row as seen in their predecessors. Morganucododontans are thought to have been insectivorous and carnivorous, their teeth adapted for shearing. Niche partitioning is known among various morganucodontan species, different types specialised for different prey. [5] Paceyodon davidi was the largest species of morganucodontans. It is known by a right lower molariform 3.3 mm (0.13 in) in length, which is bigger than molariforms of all other morganucodontans. [6]
The teeth of some morganucodonts are structured in such a way that occlusion happens by the lower cusp "a" fitting anteriorly to upper cusp "A", between "A" and "B". This occlusion pattern is by no means universal among morganucodontans, however; Dinnetherium , for example has an occlusion mechanism closer to that of the eutriconodonts, in which molars basically alternate. [7] [8] Wear facets are present.
The septomaxilla, a primitive feature also found in Sinoconodon, is found in morganucodontans, as well as a fully ossified orbitosphenoid. The anterior lamina is enlarged. The cranial moiety of the squamosal is a narrow bone that is superficially placed to the petrosal and parietal. Unlike its predecessors, the morganucodontans have a larger cerebral capacity and a longer cochlea.
The atlas elements are unfused; there is a suture between the dens and axis. The cervical ribs are not fused to the centra. The coracoid and procoracoid, which are absent in therians, are present. The head of the humerus is spherical as in mammals, but the spiral ulnar condyle is cynodont-like. In the pelvic girdle, the pubis, ilium and ischium are unfused. At least Megazostrodon and Erythrotherium are unique among basal mammaliaforms for lacking epipubic bones, suggesting that they didn't have the same reproductive constraints. [9]
Morganucodon at least appears to have had a long lifespan and a lower metabolism than modern mammals. [10]
Because morganucodontans possessed the mammalian dentary-squamosal jaw joint, systematists like G. G. Simpson (1959) [11] considered the morganucodontans to be mammals. Some palaeontologists continue to use this classification. [12] Others use crown-group terminology, which limits Mammalia to the descendants of the closest common ancestor of living mammals. By this definition, more basal orders like morganucodontans are not in Mammalia, but are mammaliaformes. [13] This is the most commonly accepted classification system.[ citation needed ]
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.
Synapsida is one of the two major clades of vertebrate animals in the group Amniota, the other being the Sauropsida. The synapsids were the dominant land animals in the late Paleozoic and early Mesozoic, but the only group that survived into the Cenozoic are mammals. 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.
Morganucodon is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, Morganucodon is well represented by abundant and well preserved material. Most of this comes from Glamorgan in Wales, but fossils have also been found in Yunnan Province in China and various parts of Europe and North America. Some closely related animals (Megazostrodon) are known from exquisite fossils from South Africa.
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, and Delsatia. The shuotheriids, another group of Jurassic mammaliaforms, also shared some dental characteristics with docodonts. One study has suggested that shuotheriids are closely related to docodonts, though others consider shuotheriids to be true mammals, perhaps related to monotremes.
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.
Sinoconodon is an extinct genus of mammaliamorphs that appears in the fossil record of the Lufeng Formation of China in the Sinemurian stage of the Early Jurassic period, about 193 million years ago. While sharing many plesiomorphic traits with other non-mammaliaform cynodonts, it possessed a special, secondarily evolved jaw joint between the dentary and the squamosal bones, which in more derived taxa would replace the primitive tetrapod one between the articular and quadrate bones. The presence of a dentary-squamosal joint is a trait historically used to define mammals.
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.
The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked like mammals. The lineage leading to today's mammals split up in the Jurassic; synapsids from this period include Dryolestes, more closely related to extant placentals and marsupials than to monotremes, as well as Ambondro, more closely related to monotremes. Later on, the eutherian and metatherian lineages separated; the metatherians are the animals more closely related to the marsupials, while the eutherians are those more closely related to the placentals. Since Juramaia, the earliest known eutherian, lived 160 million years ago in the Jurassic, this divergence must have occurred in the same period.
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.
The evolution of mammalian auditory ossicles was an evolutionary process that resulted in the formation of the bones of the mammalian middle ear. These bones, or ossicles, are a defining characteristic of all mammals. The event is well-documented and important as a demonstration of transitional forms and exaptation, the re-purposing of existing structures during evolution.
Gobiconodon is an extinct genus of carnivorous mammals belonging to the family Gobiconodontidae. Undisputed records of Gobiconodon are restricted to the Early Cretaceous of Asia and North America, but isolated teeth attributed to the genus have also been described from formations in England and Morocco dating as far back as the Middle Jurassic. Species of Gobiconodon varied considerably in size, with G. ostromi, one of the larger species, being around the size of a modern Virginia opossum. Like other gobiconodontids, it possessed several speciations towards carnivory, such as shearing molariform teeth, large canine-like incisors and powerful jaw and forelimb musculature, indicating that it probably fed on vertebrate prey. Unusually among predatory mammals and other eutriconodonts, the lower canines were vestigial, with the first lower incisor pair having become massive and canine-like. Like the larger Repenomamus there might be some evidence of scavenging.
Kuehneotherium is an early mammaliaform genus, previously considered a holothere, that lived during the Late Triassic-Early Jurassic Epochs and is characterized by reversed-triangle pattern of molar cusps. Although many fossils have been found, the fossils are limited to teeth, dental fragments, and mandible fragments. The genus includes Kuehneotherium praecursoris and all related species. It was first named and described by Doris M. Kermack, K. A. Kermack, and Frances Mussett in November 1967. The family Kuehneotheriidae and the genus Kuehneotherium were created to house the single species Kuehneotherium praecursoris. Modeling based upon a comparison of the Kuehneotherium jaw with other mammaliaforms indicates it was about the size of a modern-day shrew between 4 and 5.5 g at adulthood.
Triconodontidae is an extinct family of small, carnivorous mammals belonging to the order Eutriconodonta, endemic to what would become Asia, Europe, North America and probably also Africa and South America during the Jurassic through Cretaceous periods at least from 190–66 mya.
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.
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.
Ichthyoconodon is an extinct genus of eutriconodont mammal from the Lower Cretaceous of Morocco. It is notable for having been found in a unique marine location, and the shape of its teeth suggests an unusual, potentially fish-eating ecological niche. Analysis suggests it is part of a group of gliding mammals that includes Volaticotherium.
Borealestes is a genus of docodontan from the Middle Jurassic of Britain, first discovered on the Isle of Skye near the village of Elgol. It was the earliest mammaliaform from the Mesozoic found and named in Scotland. A second species and was later found in other Middle Jurassic sites in England, but is now shown to be a different genus. A new species, B. cuillinensis was named in 2021, also from Skye.
Wareolestes rex is a mammaliaform from the Middle Jurassic (Bathonian) rocks of England and Scotland. It was originally known from isolated teeth from England, before a more complete jaw with teeth was found in the Kilmaluag Formation of Skye, Scotland.
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.
Bocaconodon is an early mammaliaform genus that lived during the Pliensbachian of Mexico. The type and only species, Bocaconodon tamaulipensis, was named and described in 2008. It is known from a single specimen, a partial right dentary bone preserving two nearly complete molar teeth and the rear portion of a third molar. The specimen was found at the Huizachal Canyon locality, "a Pliensbachian floodplain siltstone in the La Boca Formation".
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