Megaconus Temporal range: Callovian ~ | |
---|---|
Life restoration of Megaconus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Subclass: | † Allotheria |
Genus: | † Megaconus Zhou et al., 2013 |
Type species | |
†Megaconus mammaliaformis Zhou et al., 2013 |
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. [1] 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. [2] However, later studies cast doubt on the euharamiyidan intrepretation, instead finding it to be a basal allotherian mammal. [3]
Megaconus is one of the few early mammaliaforms known from a complete skeleton. The skeleton includes both the jaw bones and the teeth, which are the most informative features because they allow for comparisons with other mammaliaforms known only from dental features. Megaconus has a dentition similar to those of rodents, with large incisors at the front of the jaws and broad molars in the back. One distinguishing feature of Megaconus is a pair of enlarged premolar teeth in the lower jaw. The teeth of Megaconus have many cusps, allowing them to interlock tightly when the jaws are closed. If Megaconus is a non-mammalian mammaliaform, it is one of the most basal mammaliaforms to possess such complex teeth. [1]
The middle ear of Megaconus is more primitive than that of modern mammals. The three bones that make up the middle ear in modern mammals — the malleus, incus, and stapes — originated from the lower jaw in the ancestors of mammals. In Megaconus, these bones are still associated with the lower jaw, placed within a groove behind the dentary bone of the lower jaw. [1]
Megaconus is estimated to have weighed about 250 grams (8.8 oz). It probably had an outwardly similar appearance to multituberculates, a major group of Mesozoic mammals. However, its body is longer than those of multituberculates and most other Mesozoic mammaliaforms, having more back vertebrae (24) than other early mammals. The transition between the thoracic and lumbar vertebrae is more gradual in Megaconus than it is in multituberculates, and the boundary between the anterior and posterior epaxial muscles (the muscles that cover the front part and the back part of the back, respectively) is positioned farther back. [1]
Megaconus is inferred to have been ambulatory, meaning that it walked on the ground. Its claws are short, meaning that they were not suitable for digging, and only slightly curved, meaning that they were not suitable for climbing. The lack of an elongated calcaneus or "heel" on the ankle means that it could not have run fast. However, Megaconus differs from most ambulatory mammals in having a fused tibia and fibula in its lower leg, a feature that is normally seen in jumping or bipedal mammals. Among the only living ambulatory mammals to have a fused tibia and fibula are the armadillo and the aardvark. [1]
Remains of fur are preserved on parts of the skeleton. The fur consists of dark guard hairs and a lighter layer of underfur. Fur seems to be absent on the underside of the abdomen. A keratinous spur on the rear leg may have been used to deliver poison, like the spur of the living platypus. [1]
The holotype skeleton of Megaconus was discovered in the Daohugou Beds of the Tiaojishan Formation in Inner Mongolia, China. The layer in which Megaconus was found is about 165 million years old (Ma). In addition to Megaconus, several other mammaliaforms are known from the Daohugou Beds: the non-mammalian mammaliaform Castorocauda , the basal mammals Volaticotherium and Pseudotribos , and the placental mammal Juramaia .
The genus name Megaconus means "large cusp", coming from the Latin mega ("large") and the Greek conus ("cusp"), in reference to the pair of large premolars in its lower jaw. The species name mammaliaformis references its ancestral mammalian features. [1]
Megaconus is part of the clade Haramiyida, which is otherwise known almost exclusively from teeth. The relationships of haramiyidans to other early mammals is contested.
One idea is that haramiyidans are close relatives of Multituberculata, the most diverse group of Mesozoic mammals, and that both are part of the larger clade Allotheria. According to this evolutionary hypothesis (phylogeny), haramiyidans fall within the crown group Mammalia — the clade originating from the most recent common ancestor of living mammals. Within the crown group Mammalia, Haramiyida would be a basal offshoot of Boreosphenida, the clade including marsupials, eutherians (placental mammals), and all mammals more closely related to them than to monotremes (which are part of the clade Australosphenida).
A second hypothesis holds that haramiyidans originated before the appearance of crown group Mammalia as more basal members of the larger clade Mammaliaformes and that multituberculates are deeply nested within the crown group, splitting up Allotheria. This phylogeny was supported by the phylogenetic analysis that included Megaconus. Features of Megaconus that link it with basal mammaliaforms include a groove in the lower jaw holding the middle ear bones (in multituberculates and other true mammals, these bones are completely separated from the lower jaw) and a calcaneum or ankle bone that lacks an enlarged heel. Below is a cladogram modified from the analysis: [1]
Mammaliaformes |
| |||||||||||||||||||||||||||||||||||||||||||||||||||
A 2022 study found it to be a basal allotherian instead, due to lacking apomorphies characteristic of euharamiyidans. [3] Furthermore, a 2020 study found it to fall outside Mammaliaformes entirely, as the sister taxon to Tritylodontidae. [4]
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.
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.
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.
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.
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.
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.
Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.
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.
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.
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.
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.