Leucocephalus Temporal range: Wuchiapingian, | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Biarmosuchia |
Family: | † Burnetiidae |
Genus: | † Leucocephalus Day et al., 2018 |
Species: | †L. wewersi |
Binomial name | |
†Leucocephalus wewersi Day et al., 2018 | |
Leucocephalus is a genus of biarmosuchian belonging to the family Burnetiidae dating to the Wuchiapingian (Late Permian). [1] It was found in the Tropidostoma Assemblage Zone of the Main Karoo Basin of South Africa. It is a monotypic taxon which contains one only species, Leucocephalus wewersi. [1] The genus name Leucocephalus is derived from Greek. Leucos, meaning white; kephalos, meaning skull, as the Leucocephalus skull discovered was unusually pale. The species epithet wewersi comes from the farm employee who found the skull, Klaus ‘Klaasie’ Wewers. [1]
Biarmosuchians are a group of some of the earliest therapsids, a group of synapsids including mammals and their ancestors. [2]
The skull of Leucocephalus was found in the Tropidostoma Assemblage Zone (Tropidostoma) of the Main Karoo Basin of South Africa. [1] Only a single skull was found which was located in 2012 at a farm called Amandelboom in Northern Cape Province. [1] It was found on a slope with strata that hosted a tetrapod fossil assemblage to the lower Tropidostoma AZ by a local sheep herder who then hung it on a fence on his farm. Although the skull was outside of its site of burial, strata stuck to it was verified to match that of the adjacent cliff section. [1]
The biostratigraphic occurrence of the skull was in the lowermost Tropidostoma Assemblage Zone. Based on previous dating of the surrounding strata, the Leucocephalus skull is estimated to be around 259 Ma. [1] Other Late Permian therapsids have been collected from the same interval and vicinity including dicynodonts (Pristerodon mackay, Tropidostoma dubium, Diictodon feliceps), a gorgonopsian, and a pareiasaur. [1] Based on discoveries of early therapsids and biarmosuchians, what is now southern Africa may have been the area of origin for burnetiamorphs. [3]
During the period Leucocephalus lived, what is considered the most extensive mass extinction in the history of the earth was occurring, [4] which caused over 80% of the all Earth's species to go extinct. [5] The cause of this end-Permian mass extinction is hypothesized to be climate change induced by volcanic CO2 degassing [6] which lead to a cascade of biotic response. [7]
Compared to their pelycosaur ancestors, Leucocephalus and other early therapsids are distinguished by more vertical (mammal like) leg positioning beneath their bodies, larger temporal fenestra and increased jaw complexity and power. [2]
Like other burnetiids, Leucocephalus skulls exhibit numerous distinguished bony protuberances and bosses giving it a bumpy appearance. These include paired supratemporal "horns" formed by the squamosals and parietals. [8] In lateral view, a ridge like boss on the nasal extending exteriorly to the prefrontal is present. [8] The snout is notably tall and the supraorbital region contains large triangular bosses. [1]
Amongst other members of the burnetiid family, Leucocephalus has some distinct features unique to its genus. This includes a longer and rounder than usual vomerine process which is visible in palatal view. The maxilla comprises the majority of the snout and is also larger than usual. Leucocephalus also has a more random arrangement of polygonal raised surfaces and irregular trenches on maxilla than what is observed in other burnetiamorphs. [1] This is hypothesized to be indicators of a thickened dermis or keratinized skin. [1] [9] Leucocephalus exhibits a sinuous intranarial process. Intranarial processes are common in mammal like therapsids, [10] however they are usually straight in biarmosuchians. [1]
A therapsid is a member of the clade Therapsida which is a major group of eupelycosaurian synapsids that includes mammals and their ancestors and 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, as opposed to the sprawling posture of many reptiles and salamanders.
Cynodonts are eutheriodont therapsids belonging to the clade Cynodontia that first appeared in the Late Permian, and extensively diversified after the Permian–Triassic extinction event. Cynodonts occupied a variety of ecological niches, both as carnivores and as herbivores. Mammals are cynodonts, as are their extinct ancestors and close relatives (Mammaliaformes), having evolved from advanced probainognathian cynodonts during the Late Triassic. All other cynodont lines went extinct, with the last known non-mammaliaform cynodont group, the Tritylodontidae, having its youngest records in the Early Cretaceous.
Diictodon is an extinct genus of pylaecephalid dicynodont. These mammal-like synapsids lived during the Late Permian period, approximately 255 million years ago. Fossils have been found in the Cistecephalus Assemblage Zone of the Madumabisa Mudstone of the Luangwa Basin in Zambia and the Tropidostoma Assemblage Zone of the Teekloof Formation, Tapinocephalus Assemblage Zone of the Abrahamskraal Formation, Dicynodon Assemblage Zone of the Balfour Formation, Cistecephalus Assemblage Zone of the Middleton or Balfour Formation of South Africa and the Guodikeng Formation of China. Roughly half of all Permian vertebrate specimens found in South Africa are those of Diictodon. This small herbivorous animal was one of the most successful synapsids in the Permian period.
Biarmosuchia is an extinct clade of non-mammalian synapsids from the Permian. Biarmosuchians are the most basal group of the therapsids. They were moderately-sized, lightly built carnivores, intermediate in form between basal sphenacodont "pelycosaurs" and more advanced therapsids. Biarmosuchians were rare components of Permian ecosystems, and the majority of species belong to the clade Burnetiamorpha, which are characterized by elaborate cranial ornamentation.
Dinogorgon is a genus of gorgonopsid from the Late Permian of South Africa and Tanzania. The generic name Dinogorgon is derived from Greek, meaning "terrible gorgon", while its species name rubidgei is taken from the surname of renowned Karoo paleontologist, Professor Bruce Rubidge, who has contributed to much of the research conducted on therapsids of the Karoo Basin. The type species of the genus is D. rubidgei.
Anteosaurus is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian during the Capitanian stage, about 265 to 260 million years ago in what is now South Africa. It is mainly known by cranial remains and few postcranial bones. With its skull reaching 80–90 cm (31–35 in) in length and a body size estimated at more than 5 m (16 ft) in length, and 500 to 600 kg in weight, Anteosaurus was the largest known carnivorous non-mammalian synapsid and the largest terrestrial predator of the Permian period. Occupying the top of the food chain in the Middle Permian, its skull, jaws and teeth show adaptations to capture large prey like the giants titanosuchids and tapinocephalids dinocephalians and large pareiasaurs.
The Tropidostoma Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the lower Teekloof Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 240 metres (790 ft), occur from east of Sutherland through to Beaufort West and Victoria West, to areas south of Graaff-Reinet. Its northernmost exposures occur west/north-west of Colesberg. The Tropidostoma Assemblage Zone is the fourth biozone of the Beaufort Group.
Burnetiidae is an extinct family of biarmosuchian therapsids that lived in the Permian period whose fossils are found in South Africa and Russia. It contains Bullacephalus, Burnetia, Mobaceras, Niuksenitia, Paraburnetia and Proburnetia.
Rubidgea is a genus of gorgonopsid from the upper Permian of South Africa and Tanzania, containing the species Rubidgea atrox. The generic name Rubidgea is sometimes believed to be derived from the surname of renowned Karoo paleontologist, Professor Bruce Rubidge, who has contributed to much of the research conducted on therapsids of the Karoo Basin. However, this generic name was actually erected in honor of Rubidge's paternal grandfather, Sydney Rubidge, who was a renowned fossil hunter. Its species name atrox is derived from Latin, meaning “fierce, savage, terrible”. Rubidgea is part of the gorgonopsian subfamily Rubidgeinae, a derived group of large-bodied gorgonopsians restricted to the Late Permian (Lopingian). The subfamily Rubidgeinae first appeared in the Tropidostoma Assemblage Zone. They reached their highest diversity in the Cistecephalus and Daptocephalus assemblage zones of the Beaufort Group in South Africa.
Lemurosaurus is a genus of extinct biarmosuchian therapsids from the Late Permian of South Africa. The generic epithet Lemursaurus is a mix of Latin, lemures “ghosts, spirits”, and Greek, sauros, “lizard”. Lemurosaurus is easily identifiable by its prominent eye crests, and large eyes. The name Lemurosaurus pricei was coined by paleontologist Robert Broom in 1949, based on a single small crushed skull, measured at approximately 86 millimeters in length, found on the Dorsfontein farm in Graaff-Reinet. To date, only two skulls of the Lemurosaurus have been discovered, so body size is unknown. The second larger, more intact, skull was found in 1974 by a team from the National Museum, Bloemfontein.
Paraburnetia is an extinct genus of biarmosuchian therapsids from the Late Permian of South Africa. It is known for its species P. sneeubergensis and belongs to the family Burnetiidae. Paraburnetia lived just before the Permian–Triassic mass extinction event.
Lobalopex is an extinct genus of biarmosuchian therapsids. It was alive during the Late Permian and has only been found in the Teekloof Formation in South Africa. The only known species of the genus is Lobalopex mordax. Lobalopex is part of the clade of Burnetiamorpha, which have fossil specimens located in multiple areas of Africa and Russia.
Bullacephalus is an extinct genus of biarmosuchian therapsids belonging to the family Burnetiidae. The type species B. jacksoni was named in 2003. It is known from a relatively complete skull and lower jaw, discovered in the Late Permian Tapinocephalus Assemblage Zone of the Beaufort Group of South Africa. This genus of therapsida lived during the Late Permian period, approximately 250 million years ago. The name Bullacephalus comes from the Latin words "bullatus," meaning "bossed" or "knobbed," and "cephalus," meaning "head." This name refers to the distinctive bony knob on the top of the therapsid's skull, which contributes to the history of this genus. This stem based taxon includes Ictidorhinus or Hippasaurs. Bullacephalus can even be characterized as having a, “skull moderately to greatly pachyostotic; swollen boss present above the postorbital bar formed by the postfrontal and postorbital; deep linear sculpturing of the snout; exclusion of the jugal from the lateral temporal fenestra”. These Therapsids have spongy bone skull roof, palatal process of premaxilla are long, diverticulum of naris adding them to the Burnetiamorph. Furthermore, the discovery of Bullacephalus has helped to refine the taxonomic classification of therapsids. Prior to its discovery, there was uncertainty regarding the relationship between different groups of therapsids, particularly the Burnetiamorpha and the Biarmosuchia. However, the distinctive features of Bullacephalus suggest that it is a member of the Burnetiamorpha, and provides a bridge between this group and the Biarmosuchia. The discovery of Bullacephalus has also highlighted the importance of continued exploration and excavation in areas that have yielded few therapsid fossils. The Beaufort Group of South Africa, where Bullacephalus was discovered, has been an important site for therapsid fossils, but much of the area remains unexplored. Further discoveries in this region and other areas around the world may provide new insights into the evolution and diversification of therapsids, as well as other groups of extinct animals. These discoveries will also help to refine our understanding of the history of life on Earth and the processes that have shaped the diversity of organisms that exist today.
Eosimops is an extinct genus of pylaecephalid dicynodonts. They were small synapsids superficially resembling modern mammals. Eosimops is known from several skull specimens, as well as one complete skeleton. Eosimops lived during the Middle Permian of South Africa.
Lycaenodon is an extinct genus of biarmosuchian therapsids from the Late Permian of South Africa. It is known from a single species, Lycaenodon longiceps, which was named by South African paleontologist Robert Broom in 1925. Both are small-bodied biarmosuchians. Two specimens are known, and both preserve only the front portions of the skull. These specimens come from the Cistecephalus Assemblage Zone of the Karoo Basin. Broom attributed the back portion of a third skull to Lycaenodon, but subsequent examiners considered it to belong to a gorgonopsian or dinocephalian and not a biarmosuchian. Most of the distinguishing features of Lycaenodon come from its palate. As a member of Biarmosuchia, the most basal group of therapsids, Lycaenodon shares many features with earlier and less mammal-like synapsids like Dimetrodon.
Raranimus is an extinct genus of therapsids of the Middle Permian. It was described in 2009 from a partial skull found in 1998 from the Dashankou locality of the Qingtoushan Formation, outcropping in the Qilian Mountains of Gansu, China. The genus is the most basal known member of the clade Therapsida, to which the later Mammalia belong.
Herpetoskylax is an extinct genus of biarmosuchians which existed in South Africa. The type species is Herpetoskylax hopsoni. It lived in the Late Permian Period.
Christian Alfred Sidor is an American vertebrate paleontologist. He is currently a Professor in the Department of Biology, University of Washington in Seattle, as well as Curator of Vertebrate Paleontology and Associate Director for Research and Collections at the Burke Museum of Natural History and Culture. His research focuses on Permian and Triassic tetrapod evolution, especially on therapsids.
The Abrahamskraal Formation is a geological formation and is found in numerous localities in the Northern Cape, Western Cape, and the Eastern Cape of South Africa. It is the lowermost formation of the Adelaide Subgroup of the Beaufort Group, a major geological group that forms part of the greater Karoo Supergroup. It represents the first fully terrestrial geological deposits of the Karoo Basin. Outcrops of the Abrahamskraal Formation are found from the small town Middelpos in its westernmost localities, then around Sutherland, the Moordenaarskaroo north of Laingsburg, Williston, Fraserburg, Leeu-Gamka, Loxton, and Victoria West in the Western Cape and Northern Cape. In the Eastern Cape outcrops are known from Rietbron, north of Klipplaat and Grahamstown, and also southwest of East London.
Lende is an extinct genus of biarmosuchian from Malawi. It contains one species, Lende chiweta, first described by Jacobs and colleagues in 2005 and is a burnetiamorph – a group of biarmosuchians characterized by numerous bosses and swellings on the skull. The type specimen was discovered in the early 1990s in the Permian Lower Bone Bed (B1) of the Chiweta Beds of Malawi, which are believed to correlate with the Cistecephalus Assemblage Zone of the South African Karoo Supergroup, the Usili Formation of Tanzania, and the Upper Madumabisa Mudstone of Zambia. The holotype of the genus Lende is MAL 290, which comprises an almost complete skull and lower jaw.