Megazostrodon Temporal range: | |
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Molariforms of Megazostrodon (A), Erythrotherium (B) and Morganucodon (C) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Mammaliaformes |
Order: | † Morganucodonta |
Family: | † Megazostrodontidae |
Genus: | † Megazostrodon Crompton & Jenkins, 1968 [1] |
Type species | |
†Megazostrodon rudnerae Crompton & Jenkins, 1968 [1] | |
Other species | |
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Megazostrodon is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old. [3] Two species are known: M. rudnerae from the Early Jurassic of Lesotho and South Africa, and M. chenali from the Late Triassic of France. [2]
The type species M. rudnerae was first discovered in 1966 in the Elliot Formation of Lesotho, southern Africa, by palaeontologist and archaeologist Ione Rudner. It was first described by A.W. Crompton and F.A. Jenkins Jr in 1968. [1] The generic name Megazostrodon means, literally, ‘large girdle tooth’ (from the Greek mega-large, zostros-girdle and don-tooth — referring to the large external cingula of the upper molars). The specific name honours Rudner for her discovery. [4]
A second species, M. chenali, was named in 2015 based on remains found in Saint-Nicolas-de-Port, France. It is named after the French palaeontologist Emmanuel Chenal. [2]
Megazostrodon was a small, shrew-like animal between 10 and 12 centimetres (3.9 and 4.7 in) long which probably ate insects and small reptiles. It is thought to have been nocturnal as it had a larger brain than earlier cynodonts and the enlarged areas of its brain were found to be those that process sounds and smells. [3] This was probably in order to avoid being in competition with the reptiles or becoming prey to the dinosaurs. [5]
Although considered a close relative of mammals, it did have some non-mammalian characteristics inherited from its predecessors: the first two vertebrae (atlas and axis) were still unfused as in earlier cynodonts, and it only had three sacral vertebrae instead of the usual mammalian five. [6] An interclavicle is also present, which is still present in monotremes but lost in the line leading to therian mammals.
Megazostrodon is the best-known genus of the family Megazostrodontidae, part of the larger group Morganucodonta. The other members of this family that are currently known are Indozostrodon , Dinnetherium , Wareolestes and Brachyzostrodon . The megazostrodontids used to be classified as members of a group of mammals called the triconodonts, which are thought to have evolved from a specific group of cynodonts [7] during the late Triassic and early Jurassic periods. However, recent classifications consider the megazostrodontids to be mammaliaforms outside of the stricter grouping of Mammalia proper, while the triconodonts remain within the mammalian crown group.
These early mammaliaforms possessed many traits which made them well-suited for an active lifestyle. They had a heterodont dentition consisting of four types of teeth: incisors, canines, premolars and molars, as opposed to the uniform (homodont) teeth of most reptiles. [8] This enabled them to chew and therefore process their food more thoroughly than their reptilian cousins. There is evidence that the movement of the mandible allowed a shearing action to chew food. [9] Their skeletons changed so that their limbs were more mobile, being less laterally splayed, [10] and allowing faster forward motion. They had a short ribcage and large lungs, which allowed efficient respiration. [8] Their lower jaw comprised a single bone — the dentary (as opposed to the multiple bones in the jaws of their ancestors, or seven different bones found in reptilian lower jaws). The other bones which once made up the jaw had reduced, and in later mammals would become incorporated into the middle ear, [3] enhancing their hearing.
Probably the most important change in the evolution of the first mammals was that their ancestors, the cynodonts, had become endothermic. This meant that they generated their own body heat, relying on the food they ate to help sustain their body temperature rather than depending on their surrounding environment. This permitted higher, more sustained activity levels during the day than reptiles (reptiles must frequently perform temperature regulation activities such as sun basking and seeking shade). It was probably the key to becoming nocturnal — a major advantage in a world where most predators were active during the day.
Like placentals and possibly Erythrotherium , Megazostrodon is unique among mammaliaforms in lacking epipubic bones. [12] It is likely that Megazostrodon, like the modern monotremes, laid eggs.
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.
Therapsida is a clade comprising a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, resulting in a more "standing" quadrupedal posture, as opposed to the lower sprawling posture of many reptiles and amphibians.
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.
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.
Eozostrodon is an extinct morganucodont mammaliaform. It lived during the Rhaetian stage of the Late Triassic. Eozostrodon is known from disarticulated teeth from South West England and estimated to have been less than 10 cm (3.9 in) in head-body length, slightly smaller than the similar-proportioned Megazostrodon.
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.
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.
Tritylodontidae is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, with several mammalian traits including erect limbs, endothermy and details of the skeleton. They were the last-known family of the non-mammaliaform synapsids, persisting into the Early Cretaceous.
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.
Prozostrodon is an extinct genus of probainognathian cynodonts that was closely related to mammals. The remains were found in Brazil and are dated to the Carnian age of the Late Triassic. The holotype has an estimated skull length of 6.7 centimetres (2.6 in), indicating that the whole animal may have been the size of a cat. The teeth were typical of advanced cynodonts, and the animal was probably a carnivore hunting reptiles and other small prey.
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.
Diarthrognathus is an extinct genus of tritheledontid cynodonts, known from fossil evidence found in South Africa and first described in 1958 by A.W. Crompton. The creature lived during the Early Jurassic period, about 200 million years ago. It was carnivorous and small, slightly smaller than Thrinaxodon, which was under 50 centimetres (20 in) long.
Haramiyida is a possibly paraphyletic 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.
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.
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.
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.
Brasilodon is an extinct genus of small, mammal-like cynodonts that lived in what is now Brazil during the Norian age of the Late Triassic epoch, about 225.42 million years ago. While no complete skeletons have been found, the length of Brasilodon has been estimated at 12 centimetres (4.7 in). Its dentition shows that it was most likely an insectivore. The genus is monotypic, containing only the species B. quadrangularis. Brasilodon belongs to the family Brasilodontidae, whose members were some of the closest relatives of mammals, the only cynodonts alive today. Two other brasilodontid genera, Brasilitherium and Minicynodon, are now considered to be junior synonyms of Brasilodon.
Lumkuia is an extinct genus of cynodont, fossils of which have been found in the Cynognathus Assemblage Zone of the Beaufort Group in the South African Karoo Basin that date back to the early Middle Triassic. It contains a single species, Lumkuia fuzzi, which was named in 2001 on the basis of the holotype specimen BP/1/2669, which can now be found at the Bernard Price Institute in Johannesburg, South Africa. The genus has been placed in its own family, Lumkuiidae. Lumkuia is not as common as other cynodonts from the same locality such as Diademodon and Trirachodon.
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.
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.