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''Leucocephalus'' is a genus of
biarmosuchia 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 " pel ...
n belonging to the family
Burnetiidae Burnetiidae is an extinct family of biarmosuchian therapsids that lived in the Permian period whose fossils are found in South Africa, Zambia and Russia. It contains ''Bondoceras'', ''Bullacephalus'', ''Burnetia'', ''Mobaceras'', ''Niuksenitia'' ...
dating to the
Wuchiapingian In the geologic timescale, the Wuchiapingian or Wujiapingian (from in the Liangshan area of Hanzhong, Shaanxi Province ) is an age or stage of the Permian. It is also the lower or earlier of two subdivisions of the Lopingian Epoch or S ...
(
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
). It was found in the ''Tropidostoma'' Assemblage Zone of the Main Karoo Basin of
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. It is a
monotypic taxon In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of Genus, genera, the term ...
which contains one only species, ''Leucocephalus wewersi''. 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.Michael O. Day; Roger M. H. Smith; Julien Benoit; Vincent Fernandez; Bruce S. Rubidge (2018). "A new species of burnetiid (Therapsida, Burnetiamorpha) from the early Wuchiapingian of South Africa and implications for the evolutionary ecology of the family Burnetiidae". Papers in Palaeontology. Online edition. .
Biarmosuchia 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 " pel ...
ns are a group of some of the earliest therapsids, a group of
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
s including
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s and their ancestors.


Discovery

The skull of ''Leucocephalus'' was found in the
Tropidostoma Assemblage Zone 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 Stratigraphic unit, Gro ...
(Tropidostoma) of the Main Karoo Basin of South Africa. Only a single skull was found which was located in 2012 at a farm called Amandelboom in
Northern Cape Province The Northern Cape ( ; ; ) is the largest and most sparsely populated province of South Africa. It was created in 1994 when the Cape Province was split up. Its capital is Kimberley. It includes the Kalahari Gemsbok National Park, part of the Kga ...
. It was found on a slope with
strata In geology and related fields, a stratum (: strata) is a layer of Rock (geology), rock or sediment characterized by certain Lithology, lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by v ...
that hosted a tetrapod fossil assemblage to the lower ''Tropidostoma'' AZ by a local
sheep herder A shepherd is a person who tends, herds, feeds, or guards flocks of sheep. Shepherding is one of the world's oldest occupations; it exists in many parts of the globe, and it is an important part of Pastoralism, pastoralist animal husbandry. ...
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.


Paleoenvironment

The
biostratigraphic Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. "Biostratigraphy." ''Oxford Reference: Dictionary of Biology ...
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. 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. Based on discoveries of early therapsids and biarmosuchians, what is now southern Africa may have been the area of origin for burnetiamorphs. During the period ''Leucocephalus'' lived, what is considered the most extensive
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp fall in the diversity and abundance of multicellular organisms. It occ ...
in the history of the earth was occurring, which caused over 80% of the all Earth's species to go extinct. The cause of this end-Permian mass extinction is hypothesized to be climate change induced by volcanic
degassing Degassing, also known as degasification, is the removal of dissolved gases from liquids, especially water or aqueous solutions. There are numerous methods for removing gases from liquids. Gases are removed for various reasons. Chemists remove gas ...
which lead to a cascade of biotic response.


Description and paleobiology

Compared to their
pelycosaur Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now thoug ...
ancestors, ''Leucocephalus'' and other early therapsids are distinguished by more vertical (mammal like) leg positioning beneath their bodies, larger
temporal fenestra Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
and increased jaw complexity and power. 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. In lateral view, a ridge like boss on the nasal extending exteriorly to the prefrontal is present.Rubidge, Bruce & Sidor, Christian. (2002). On the cranial morphology of the basal therapsids ''Burnetia'' and ''Proburnetia'' (Therapsida: Burnetiidae). Journal of Vertebrate Paleontology. 22. 257-267. 10.1671/0272-4634(2002)022 257:OTCMOT.0.CO;2. The snout is notably tall and the supraorbital region contains large triangular bosses. 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. This is hypothesized to be indicators of a thickened dermis or
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
ized skin. ''Leucocephalus'' exhibits a sinuous intranarial process. Intranarial processes are common in mammal like therapsids, however they are usually straight in biarmosuchians.


References

{{Taxonbar, from=Q55346359, from2=Q55360243 Burnetiamorpha Prehistoric therapsid genera Wuchiapingian genera Lopingian synapsids of Africa Permian South Africa Fossils of South Africa Fossil taxa described in 2018 Lopingian life Wuchiapingian life