Megalocephalus Temporal range: Late Carboniferous, | |
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Skull diagram of Megalocephalus pachycephalus in dorsal view (left) and ventral view (right) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Sarcopterygii |
Clade: | Tetrapodomorpha |
Family: | † Baphetidae |
Subfamily: | † Loxommatinae |
Genus: | † Megalocephalus Barkas, 1873 |
Type species | |
†Orthosaurus pachycephalus Barkas, 1873 | |
Species | |
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Synonyms | |
Synonyms of M. pachycephalus
Synonyms of M. lineolatus
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Megalocephalus (meaning "big head") is an extinct genus of baphetid tetrapodomorph from the late Carboniferous (Westphalian A-C) of the British Isles and the United States (Ohio). [1] It contains two species, M. pachycephalus and M. lineolatus. [2]
Megalocephalus pachycephalus is one of the most common large tetrapodomorphs from the Carboniferous of Britain, and is known from several sites in England, Scotland, and Ireland. The first known fossils of the genus were cranial fragments which were misattributed to Rhizodus jaw bones upon their discovery in 1864. These fragments, and many subsequent discoveries, were recovered from a quarry near Newsham, Northumberland. Other Megalocephalus fossils found in the 1860s had been referred to other large amphibians, namely Pteroplax and Anthracosaurus . Two well-preserved skulls and associated jaw bones were found in Newsham in 1870 and 1871, though these were also mistakenly referred to a pre-existing genus, Loxomma . In 1873, Thomas P. Barkas named a pair of new genera for two more Newsham skulls. One was described as a supposed new reptile, Orthosaurus pachycephalus, while the other was given the name Megalocephalus macromma. [3]
Megalocephalus fossils continued to be collected in the coming decades, though most were initially referred to Loxomma. "Orthosaurus" was re-established as a distinct genus of loxommatid amphibian by D.M.S. Watson in 1926. However, it was later determined to be a preoccupied name, and the name Orthosauriscus pachycephalus was invented and used for the rest of the early 20th century. In 1947, Alfred Romer decided to rename Orthosauriscus pachycephalus to Megalocephalus brevicornis, combining Barkas' 1873 genus with "Pteroplax" brevicornis, a species named for Megalocephalus fossils in 1870. However, the "Pteroplax" brevicornis fossils were poorly described and probably destroyed by a 1909 fire, so few subsequent paleontologists use Romer's combination. The modern combination Megalocephalus pachycephalus was popularized by Eileen H. Beamont's 1977 redescription of available "loxommatid" fossils. [3]
The second apparently valid species in the genus is Megalocephalus lineolatus. This species is from the fossiliferous coal mine of Linton, Ohio, and was originally named as "Leptophractus" lineolatus by E.D. Cope in 1877. In the 1920s, Watson placed the species within "Orthosaurus", based on both Cope's specimens and new loxommatid fossils from Linton. Romer referred all Linton loxommatid material to Macrerpeton huxleyi . However, later investigation revealed that Romer's conception of Macrerpeton was chimeric, including an edopoid temnospondyl (Macrerpeton), a loxommatid (referred to Baphetes ), and possibly several more species of loxommatids. Another skull described in 1957 demonstrated that "Leptophractus" lineolatus was a species of Megalocephalus, as it showed several diagnostic traits of the genus. [3] Megalocephalus lineolatus has also been mentioned under the name "Megalocephalus enchodus", based on a belief that the lineolatus species name should instead be assigned to the Baphetes material. [4]
Megalocephalus was an amphibian, measuring 1.5 metres (4.9 ft) from head to tail. [1]
It is only known from its skull, like most other baphetids. The skull of Megalocephalus was 30 centimetres (12 in) long. The teeth of Megalocephalus were long and pointed. Baphetids had a small elongation of their eye sockets, the current use of which is not certain. It has been suggested that the elongation was housing for a salt gland to get rid of excess salt, or an extra region for jaw muscle attachments, to allow a harder bite force. [1]
Like Kyrinion , the lacrimal is excluded from the nares (nostrils) and the dentary has large fangs. [5] A 2009 phylogenetic analysis suggests that Megalocephalus is paraphyletic relative to Kyrinion, with M. pachycephalus closer to K. martilli than to M. lineolatus. Likewise, Loxomma forms a paraphyletic grade ancestral to Megalocephalus. [6]
A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods, such as snakes, legless lizards, and caecilians, have evolved to become limbless via mutations of the Hox gene. Nevertheless, these limbless groups still qualify as tetrapods through their ancestry, and some retain a pair of vestigial spurs that are remnants of the hindlimbs.
Baphetidae is an extinct family of stem-tetrapods. Baphetids were large labyrinthodont predators of the Late Carboniferous period of Europe. Fragmentary remains from the Early Carboniferous of Canada have been tentatively assigned to the group. The phylogenetic relationships of baphetids is uncertain; while many studies have placed the group as a close relative of Amniota, other analyses have found Baphetidae to be a more basal clade of early stem tetrapods. Baphetids were among the first of the Carboniferous fossil tetrapods to be found and were originally described in 1850 by John William Dawson. The baphetids have been referred to the family Loxommatidae, but this group was later shown to be a junior synonym of Baphetidae, which was named earlier in 1865. Baphetids are known mainly from skulls; very little postcranial material has been found.
Sarcopterygii — sometimes considered synonymous with Crossopterygii — is a clade of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates are characterised by prominent muscular limb buds (lobes) within their fins, which are supported by articulated appendicular skeletons. This is in contrast to the other clade of bony fish, the Actinopterygii, which have only skin-covered bony spines supporting the fins.
Ichthyostega is an extinct genus of limbed tetrapodomorphs from the Late Devonian of what is now Greenland. It was among the earliest four-limbed vertebrates ever in the fossil record and was one of the first with weight-bearing adaptations for terrestrial locomotion. Ichthyostega possessed lungs and limbs that helped it navigate through shallow water in swamps. Although Ichthyostega is often labelled a 'tetrapod' because of its limbs and fingers, it evolved long before true crown group tetrapods and could more accurately be referred to as a stegocephalian or stem tetrapod. Likewise, while undoubtedly of amphibian build and habit, it is not a true member of the group in the narrow sense, as the first modern amphibians appeared in the Triassic Period. Until finds of other early stegocephalians and closely related fishes in the late 20th century, Ichthyostega stood alone as a transitional fossil between fish and tetrapods, combining fish and tetrapod features. Newer research has shown that it had an unusual anatomy, functioning more akin to a seal than a salamander, as previously assumed.
"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.
Eucritta is an extinct genus of stem-tetrapod from the Viséan epoch in the Carboniferous period of Scotland. The name of the type and only species, E. melanolimnetes is a homage to the 1954 horror film Creature from the Black Lagoon.
Nectridea is the name of an extinct order of lepospondyl tetrapods from the Carboniferous and Permian periods, including animals such as Diplocaulus. In appearance, they would have resembled modern newts or aquatic salamanders, although they are not close relatives of modern amphibians. They were characterized by long, flattened tails to aid in swimming, as well as numerous features of the vertebrae.
Pholiderpeton is an extinct genus of embolomere amphibian which lived in the Late Carboniferous period (Bashkirian) of England. The genus was first named by Thomas Henry Huxley in 1869 to include the species P. scutigerum, based on the disarticulated front half of a skeleton discovered near Bradford, Yorkshire. Associated fossil wood suggests that this specimen died inside a Lepidodendron tree trunk.
Diplovertebron is an extinct genus of embolomere that lived in the Late Carboniferous period (Moscovian), about 310 million years ago. Diplovertebron was a medium-sized animal, around 50 cm in length. Members of the genus inhabited European Carboniferous swamps in what is now the Czech Republic. They were closely related to larger swamp-dwelling tetrapods like Proterogyrinus and Anthracosaurus. However, Diplovertebron were much smaller than these large, crocodile-like creatures. Known from a single species, Diplovertebron punctatum, this genus has had a complicated history closely tied to Gephyrostegus, another genus of small, reptile-like amphibians.
Anthracosaurus is an extinct genus of embolomere that lived during the Late Carboniferous in what is now Scotland, England, and Ohio. Measuring around 10 ft (3.0 m) long, it was a large, aquatic eel-like predator. It has a robust skull about 40 centimetres (1.3 ft) in length with large teeth in the jaws and on the roof of the mouth. Anthracosaurus probably inhabited swamps, rivers and lakes. Its name is Greek for "coal lizard".
Pederpes is an extinct genus of early Carboniferous stegocephalian, dating from 348 to 347.6 Ma in the Tournaisian age. Pederpes contains one species, P. finneyae, 1 m long.
Spathicephalus is an extinct genus of stem tetrapods that lived during the middle of the Carboniferous Period. The genus includes two species: the type species S. mirus from Scotland, which is known from two mostly complete skulls and other cranial material, and the species S. pereger from Nova Scotia, which is known from a single fragment of the skull table. Based on the S. mirus material, the appearance of Spathicephalus is unlike that of any other early tetrapod, with a flattened, square-shaped skull and jaws lined with hundreds of very small chisel-like teeth. However, Spathicephalus shares several anatomical features with a family of stem tetrapods called Baphetidae, leading most paleontologists who have studied the genus to place it within a larger group called Baphetoidea, often as part of its own monotypic family Spathicephalidae. Spathicephalus is thought to have fed on aquatic invertebrates through a combination of suction feeding and filter feeding.
Loxomma is an extinct genus of Loxommatinae and one of the first Carboniferous tetrapods. They were first described in 1862 and further described in 1870 when two more craniums were found. It is mostly associated with the area of the United Kingdom. They share features with modern reptiles as well as with fish. They had 4 paddle-like limbs that they used to swim in lakes, but they breathed air. Their diet consisted mostly of live fish. They are of the family Baphetidae which are distinguished by their keyhole shaped orbits, while Loxomma themselves are distinguished by the unique texture on their skulls, said to be honeycomb-like.
Neopteroplax is an extinct genus of eogyrinid embolomere closely related to European genera such as Eogyrinus and Pteroplax. Members of this genus were among the largest embolomeres in North America. Neopteroplax is primarily known from a large skull found in Ohio, although fragmentary embolomere fossils from Texas and New Mexico have also been tentatively referred to the genus. Despite its similarities to specific European embolomeres, it can be distinguished from them due to a small number of skull and jaw features, most notably a lower surangular at the upper rear portion of the lower jaw.
Odonterpeton is an extinct genus of "microsaur" from the Late Carboniferous of Ohio, containing the lone species Odonterpeton triangulare. It is known from a single partial skeleton preserving the skull, forelimbs, and the front part of the torso. The specimen was found in the abandoned Diamond Coal Mine of Linton, Ohio, a fossiliferous coal deposit dated to the late Moscovian stage, about 310 million years ago.
Kyrinion is an extinct genus of baphetid tetrapodomorph from the Late Carboniferous of England. It is known from a skull that was found in Tyne and Wear county dating back to the Westphalian stage. Along with the skull is part of the lower jaw, an arch of the atlas bone and a rib possibly belonging to a cervical (neck) vertebra. The type species K. martilli was named from this material in 2003.
Adamanterpeton is a genus of Edopoid Temnospondyl within the family Cochleosauridae. The type species A. ohioensis was named in 1998 and is currently the only known species within this genus. Adamanterpeton is rare in the Linton vertebrate assemblage, with other amphibians like Sauropleura, Ophiderpeton, and Colosteus being more common. Unlike other Linton vertebrates, Adamanterpeton may have been adapted to a terrestrial lifestyle.
Occidens is an extinct genus of stem tetrapod from the Early Carboniferous (Tournaisian) Altagoan Formation of Northern Ireland. It is known from a single type species, Occidens portlocki, named in 2004 on the basis of a left lower jaw described by British geologist Joseph Ellison Portlock in 1843.
Eobaphetes is an extinct genus of embolomere which likely lived in the Pennsylvanian of Kansas. The genus is based on several skull and jaw fragments of a single individual. They were originally described under the species Erpetosuchus kansasensis, but this was later changed to Eobaphetes kansasensis when it was determined that Erpetosuchus was preoccupied by a Triassic reptile.
Diploradus is an extinct genus of four-limbed stem-tetrapod from the Mississippian (mid-Tournaisian) of Scotland. It contains a single species, Diploradus austiumensis, based on an incomplete skull and jaw fragments from the Ballagan Formation at Burnmouth. The most complete part of the specimen, the lower jaw, was about 3.0 cm in length and possessed several rows of small, numerous teeth. It likely represents a juvenile animal. Diploradus was described in a 2016 study which was devised to fill in the tetrapod and stem-tetrapod faunas of Romer's gap, an interval of the early Carboniferous with few vertebrate fossils. It was one of five new genera named in this study, along with Aytonerpeton, Koilops, Ossirarus, and Perittodus.