Styracocephalus

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Styracocephalus
Temporal range: Capitanian, 265.8–260.4  Ma
StyracocephalusDB.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Suborder: Dinocephalia
Family: Styracocephalidae
Genus: Styracocephalus
Haughton, 1929
Species:
S. platyrhynchus
Binomial name
Styracocephalus platyrhynchus
Haughton, 1929
Styracocephalus STYRACOCEPHALUS 2 copia.jpg
Styracocephalus

Styracocephalus platyrhynchus (Greek for "spiked-head") is an extinct genus of dinocephalian therapsid that existed during the mid-Permian throughout South Africa, but mainly in the Karoo Basin. It is often referred to by its single known species Styracocephalus platyrhynchus. The Dinocephalia clade consisted of the largest land vertebrates and herbivores during the early to mid-Permian. This period is often also referred to as the Guadalupian epoch, approximately 270 to 260 million years ago. [1] [2]

Contents

Styracocephalus Skull from Right Lateral View, Specimen BP/1/7141, found in the Sutherland District of South Africa Styracocephalus specimen BP17141 from Lateral View.png
Styracocephalus Skull from Right Lateral View, Specimen BP/1/7141, found in the Sutherland District of South Africa

Although multiple Styracocephalus skulls have been recovered, there has yet to be a specimen found of the entire therapsid skeleton. A majority of the skulls collected have also been mature skulls, as the juvenile tapinocephalid skulls are identified by having a small non-fused basioccipital. [3] The presence of enlarged canines, pachyostotic cranial boss, and horns are all plesiomorphic traits found in Dinocephalia. [2] Styracocephalus's head ornament meant that it could be recognised from a distance. One of the most striking feature of Styracocephalus is the large backward-protruding tabular horns. [4] It was around 1.8 metres (5 ft 11 in) in length, [5] with a 42-centimetre-long (17 in), 29-centimetre-wide (11 in) skull. [4]

History of Discovery

The skull of the holotype of Styracocephalus platyrhynchus was found in 1928 by L.D. Boonstra on the farm Boesmansivier, South Africa. Skull of Holotype of Styracocephalus SAM PK8936.png
The skull of the holotype of Styracocephalus platyrhynchus was found in 1928 by L.D. Boonstra on the farm Boesmansivier, South Africa.

The first Styracocephalus fossil was discovered by Lieuwe Dirk Boonstra in 1928 from a Tapinocephalus bed on a farm called Boesmans Rivier, in the Beaufort West Division. [1] The original skeleton was crushed, but still showed some unique features such as outward projecting tabular horns, a shallow snout, and a small temporal opening. The skull had a maximum length of around 400 mm and a width of 390 mm. [4] Upon initial discovery, Boonstra compared the newfound specimen to Burnetia stating that the palate was more like that of a gorgonopsian than a therapsid. When Boonstra evaluated its place as a therapsid, it was found to contain characteristics resembling Therocephalia, Gorgonopsia, as well as Dinocephalia. [6] The first classification of Styracocephalus was in 1929 by S. H. Haughton and he placed it in its own new subclade of Therapsida due to its unique blend of features. [1] This original holotype that was found is called SAM 8936. [6]

Description

Skull

Although the SAM 8936 holotype was used to identify many characteristics of the skull, more recent specimens such as the SAM K 8071 specimen helped determine the posterior of the skull. The SAM K 8071 specimen was twice the size of the original holotype showing the variety seen in the morphological size of Styracocephalus. [6]

A composite diagram of Styracocephalus skull from the lateral view Composite diagram of Styracocephalus skull in Lateral View.png
A composite diagram of Styracocephalus skull from the lateral view

Styracocephalus has a narrow and long snout, with thickened postorbital bones. The median nasal boss is convex and is not connected to the extremely pachyostosed interorbital section. The cranial pachyostosis is split into four regions, the medial nasal boss, a think interorbital skull roof, paired posterior postorbital horns, and the squamosal boss that flares out laterally. [1] The frontal bone on the skull roof is not part of the dorsal rim of the orbit and instead extends anteriorly between the parietal and the nasal on the skull roof. The postfrontal is large and makes contact with both the frontal and the post-orbital. The tabular found on the dorsal lateral of the occiput of Styracocephalus is roughly rectangular and has been known to be variable in size. The postorbital includes a significant amount of the boss above the orbital, as well as the dorsal surface of the horn. It is in junction with the post-frontal, parietal, and squamosal. There are two parietals that form a pair on the midline in a triangular shape, also containing a small pineal foramen. [4] The occiput is rectangularly shaped and has thickened squamosal crests and these crests extend ventrally to the single temporal fenestrae. It also has a small temporal opening with the postorbital extending above it anteriorly. [6] A rather significant characteristic of the skull would be the tabular horns that extend laterally backward. The horn is one of the most distinguishable traits of Styracocephalus, and the name "spike-head" refers to these curved horns. The stapes in the skull are short with the distal end slightly swollen. [4] Most of the basicranial elements on known specimens of Strycacocephalus are poorly preserved; however, some portions such as the stapes are uniquely distinguishable and appear as small dumbbell-shaped bones that contact the fenestra and quadrate. When the skull is viewed posteriorly, it takes on a more square-like configuration. [1] [6]

Dentition

Segmented CT scan reconstruction of the anterior fragment of the left lower jaw of SAM-PK-8936, with the arrow pointing at talon-heel dinocephalian morphology Segmented CT scan reconstruction of the anterior fragment of the left lower jaw of Styracocephalus.png
Segmented CT scan reconstruction of the anterior fragment of the left lower jaw of SAM-PK-8936, with the arrow pointing at talon–heel dinocephalian morphology

Styracocephalus has large non-serrated canines which is not typical of tapinocephalids, except for Tapinocaninus and Ulemosaurus . [1] [4] They also have heeled incisors and pronounced canines on the upper and lower jaw; however, Styracocephalus specifically has bulbous post-canines whereas other tapinocephalids often have leaf-shaped post-canines. [2]

There are typically around eight to ten of these post-canines that also have lingual heels. The incisors containing crushing heels are like those seen in Tapinocephalidae, Titanosuchidae and Dinocephalia. The presence of lingual heels indicates the specimen is representative of Dinocephalia later in the mid-Permian. They also have crowns that are relatively blunt, which can be indicative of grinding plant material. [6]

Lower Jaw

Composite diagram of Styracocephalus skull from ventral view, depicting the palate Composite diagrams of Styracocephalus skull in Ventral View.png
Composite diagram of Styracocephalus skull from ventral view, depicting the palate

The lower jaw of Styracocephalus has a large dentary that occupies almost three-fourths of the jaw when viewed laterally. The anterior margin of the jaw slopes posteroventral forming a slight chin on the anterior side of the lower jaw. The angular makes up a broad surface posterior to the dentary and forms a junction with the prearticular. The splenial is flat on the medial side of the jaw and resembles the shape of a spindle. The coronoid is elongated on the lower jaw immediately behind the canines, and also makes contact with the dentary. [1] [4] [6]

Palate

The palate of Styracocephalus is often well preserved in many specimens. It has two paired pterygoids that make up a flat triangular surface that contacts and is concave to the dentary. The interpterygoidal vacuity is small and, on the midline, behind the transverse process. [6] There are small palatal teeth as well as paired vomers on the midline. Serrations, although not found on Styracocephalus, are found on Titanophoneus and Theriodontia. The palate also does not exhibit any suborbital fenestrae. [1]

Paleobiology

The Tapinocephalidae were one of the first earliest groups of herbivores as seen by their talon and heel-like dentition. The enamel on the heel of the dentition for many tapinocephalid specimens has signs of reduced deposition indicating the grinding action of consuming plant material. [2] Due to the smaller body size of Styracocephalus, it likely consumed smaller vegetative plants. Based on dinocephalian dentition, the presence of a large, heavy cranium and a poorly attached fragile mandible makes it difficult for this species to consume tough vegetation on dry ground. [7] One proposed purpose of horns on dinocephalians is to adapt to a less individualized behavioral pattern. With cranial ornaments especially, horns, that are often used for headbutting, this species demonstrates a more social and group-like behavior. [2]

Paleoecology

Where the Styracocephalus skull of BP/1/7141 locality is on Tontelbosch, visualized by the red dot. Map of South Africa showing the locality in the Sutherland district, Northern Cape Province, with picture depicting area where Styracocephalus fossils are often found Map of South Africa showing the locality in the Sutherland district, Northern Cape Province.png
Where the Styracocephalus skull of BP/1/7141 locality is on Tontelbosch, visualized by the red dot. Map of South Africa showing the locality in the Sutherland district, Northern Cape Province, with picture depicting area where Styracocephalus fossils are often found

The majority of dinocephalian fossils such as that of Styracocephalus are found in the Tapinocephalus Assemblage Zone of the South African Karoo. [1] [7] Many of the more advanced tapinocephalid fossils, which exhibit characteristics seen in later-mid-Permian Dinocephalia, are often found in this assemblage zone. [3] However, recent recoveries of Mid-Permian Dinocephalia have been in South America, suggesting that they were dispersed in various locations across Pangea. This also supports that Therapsida were found on a global scale from as early as the Guadalupian including Eastern Europe and South America. [8] According to research conducted on the Karoo Basin in South Africa, Styracocephalus likely preferred warmer climate conditions with fluctuating precipitation. The basin was formed during Pangea time and survived through the breaking up of the continent, which likely contributes to the thorough global presence of Dinocephalia. [9] Many tapinocephalid fossils have also been discovered on farms such as the holotype found on the Boesmans Rivier farm, recent dinocephalian fossils have also been found on other farms like the Klein Wolwefontein. Consistent with assumptions made by Boonstra on characteristics of Dinocephalia, it can be inferred that Styracocephalus spent a fair amount of time in shallow ponds and marshlands. The ratio between the cranial elements and limbs of this group of Dinocephalia meant a decreased likelihood of long land travel, and a much higher probability of dwelling in marsh areas and eating nearby marsh vegetation. [7] [10]

Classification

The Styracocephalus placement has long been questioned in the past due to debates regarding the familial classifications of Dinocephalia. It was previously thought that Styracocephalus made up its own dinocephalian family, as with Anterosauridae, Titanosuchidae, and Tapinocephalidae. The two reduced modern dinocephalian family subclade classifications are Anteosauria and Tapinocephalia, with Styracocephalidae and sister taxa Estemmenosuchidae classifying as tapinocephalids. Below is a cladogram depicting the relationship of the Styracocephalidae with other dinocephalians based on a phylogenetic study published in 2019. [1] [11]

Therapsida

Dimetrodon

Dinocephalia
Tapinocephalia

Estemmenosuchus

Styracocephalus

Jonkeria

Ulemosaurus

Moschops

Anteosauridae

Gorgonopsia

Anomodontia

Biarmosuchia

Biarmosuchus

Hipposaurus

Herpetoskylax

Lemurosaurus

Proburnetia

Burnetia

Related Research Articles

<span class="mw-page-title-main">Dinocephalia</span> Extinct clade of stem-mammals

Dinocephalians are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. Dinocephalians included herbivorous, carnivorous, and omnivorous forms. Many species had thickened skulls with many knobs and bony projections. Dinocephalians were the first non-mammalian therapsids to be scientifically described and their fossils are known from Russia, China, Brazil, South Africa, Zimbabwe, and Tanzania.

<i>Moschops</i> Extinct genus of therapsids that lived in the Guadalupian epoch

Moschops is an extinct genus of therapsids that lived in the Guadalupian epoch, around 265–260 million years ago. They were heavily built plant eaters, and they may have lived partly in water, as hippopotamuses do. They had short, thick heads and might have competed by head-butting each other. Their elbow joints allowed them to walk with a more mammal-like gait rather than crawling. Their remains were found in the Karoo region of South Africa, belonging to the Tapinocephalus Assemblage Zone. Therapsids, such as Moschops, are synapsids, the dominant land animals in the Permian period, which ended 252 million years ago.

<span class="mw-page-title-main">Anteosaur</span> Extinct clade of therapsids

Anteosaurs are a group of large, primitive carnivorous dinocephalian therapsids with large canines and incisors and short limbs, that are known from the Middle Permian of South Africa, Russia, China, and Brazil. Some grew very large, with skulls 50–80 centimetres (20–31 in) long, and were the largest predators of their time. They died out at the end of the Middle Permian, possibly as a result of the extinction of the herbivorous Tapinocephalia on which they may have fed.

<span class="mw-page-title-main">Anteosauridae</span> Extinct family of therapsids

Anteosauridae is an extinct family of large carnivorous dinocephalian therapsids that are known from the Middle Permian of Asia, Africa, and South America.These animals were by far the largest predators of the Permian period, with skulls reaching 80 cm in length in adult individuals, far larger than the biggest gorgonopsian.

<i>Anteosaurus</i> Extinct genus of anteosaurid synapsid from the Permian

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. Measuring 5–6 m (16–20 ft) long and weighing about 600 kg (1,300 lb), 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.

<i>Tapinocephalus</i> Assemblage Zone

The Tapinocephalus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the middle Abrahamskraal 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 2,000 metres (6,600 ft), occur from Merweville and Leeu-Gamka in its southernmost exposures, from Sutherland through to Beaufort West where outcrops start to only be found in the south-east, north of Oudshoorn and Willowmore, reaching up to areas south of Graaff-Reinet. Its northernmost exposures occur around the towns Fraserburg and Victoria West. The Tapinocephalus Assemblage Zone is the second biozone of the Beaufort Group.

<i>Struthiocephalus</i> Extinct genus of therapsids

Struthiocephalus is an extinct genus of dinocephalian therapsids from the Permian of South Africa. It was a large animal, reaching 288 kg (635 lb) in body mass.

<i>Tapinocaninus</i> Extinct genus of therapsids

Tapinocaninus is an extinct genus of therapsids in the family Tapinocephalidae, of which it is the most basal member. Only one species is known, Tapinocaninus pamelae. The species is named in honor of Rubidge's mother, Pam. Fossils have been found dating from the Middle Permian.

<i>Theriognathus</i> Extinct genus of therapsids from late Permian South Africa and Tanzania

Theriognathus is an extinct genus of therocephalian therapsid belonging to the family Whaitsiidae, known from fossils from South Africa, Zambia, and Tanzania. Theriognathus has been dated as existing during the Late Permian. Although Theriognathus means mammal jaw, the lower jaw is actually made up of several bones as seen in modern reptiles, in contrast to mammals. Theriognathus displayed many different reptilian and mammalian characteristics. For example, Theriognathus had canine teeth like mammals, and a secondary palate, multiple bones in the mandible, and a typical reptilian jaw joint, all characteristics of reptiles. It is speculated that Theriognathus was either carnivorous or omnivorous based on its teeth, and was suited to hunting small prey in undergrowth. This synapsid adopted a sleek profile of a mammalian predator, with a narrow snout and around 1 meter long. Theriognathus is represented by 56 specimens in the fossil record.

<i>Aelurosaurus</i> Extinct genus of therapsids

Aelurosaurus is a small, carnivorous, extinct genus of gorgonopsian therapsids from the Late Permian of South Africa. It was discovered in the Karoo Basin of South Africa, and first named by Richard Owen in 1881. It was named so because it appeared to be an ancestor for cat-like marsupials, but not yet a mammal itself. It contains five species, A. felinus, A. whaitsi, A. polyodon, A. wilmanae, and A.? watermeyeri. A. felinus, the type species, is generally well described with established features, while the other four species are not due to their poorly preserved holotypes.

<i>Biseridens</i> Extinct genus of therapsids

Biseridens is an extinct genus of anomodont therapsid, and one of the most basal anomodont genera known. Originally known from a partial skull misidentified as an eotitanosuchian in 1997, another well-preserved skull was found in the Qingtoushan Formation in the Qilian Mountains of Gansu, China, in 2009 that clarified its relationships to anomodonts, such as the dicynodonts.

<i>Moschognathus</i> Extinct genus of therapsids that lived in the Guadalupian epoch

Moschognathus is an extinct genus of dinocephalian therapsid in the family Tapinocephalidae. The genus includes only the type species M. whaitsi, named by palaeontologist Robert Broom in 1914. It was a short-snouted tapinocephalid, closely related to and resembling the well-known genus Moschops, but its skull is less thickened overall has a relatively longer and shallower snout by comparison. Indeed, Moschognathus has typically been regarded as a junior synonym of Moschops since 1969 after Lieuwe Dirk Boonstra sunk Moschognathus into Moschops, albeit retained as its own doubtfully valid species. However, researchers in the 21st century have expressed doubt over this synonymy and suggested that Moschognathus is a distinct taxon after all, including first by Christian Kammerer in a 2009 Ph.D. thesis and formally in 2015 by Alessandra D. S. Boos and colleagues in 2015. Moschognathus has since began to re-enter scientific literature of dinocephalians as a valid name and treated distinct from Moschops.

<i>Pachydectes</i> Extinct genus of therapsids

Pachydectes is an extinct genus of biarmosuchian therapsids from the Middle Permian of South Africa known from a single skull. The etymology of the name Pachydectes is derived from the Greek word pakhus, meaning "thick" or "thickened", and dektes, meaning "biter". In conjunction this name is representative of the unique pachyostotic bone present above the maxillary canine tooth found in the skull of the specimen. There is only one known species within the genus, Pachydectes elsi which is named in honor of the person who discovered the fossil.

<i>Scymnosaurus</i> Extinct genus of therapsids from middle Permian South Africa

Scymnosaurus is a dubious genus of therocephalian therapsids based upon various fossils of large early therocephalians. The genus was described by Robert Broom in 1903 with S. ferox, followed by S. watsoni in 1915 and a third, S. major, by Lieuwe Dirk Boonstra in 1954. Each of these species are considered nomen dubia today and based upon specimens belonging to two separate families of therocephalians. S. ferox and S. major represent specimens of Lycosuchidae incertae sedis, while S. watsoni is Scylacosauridae incertae sedis. Broom named a fourth species in 1907 from KwaZulu-Natal, S. warreni, though he later referred it to Moschorhinus as a valid species in 1932 but now is recognised as being synonymous with M. kitchingi.

<i>Criocephalosaurus</i> Extinct genus of therapsids

Criocephalosaurus is an extinct genus of tapinocephalian therapsids that lived in Southern Africa during the Guadalupian epoch of the Permian. They are the latest surviving dinocephalians, extending past the Abrahamskraal Formation into the lowermost Poortjie Member of the Teekloof Formation in South Africa. They are also regarded as the most derived of the dinocephalians, alongside Tapinocephalus, and the most abundant in the fossil record.

<span class="mw-page-title-main">Abrahamskraal Formation</span> Geological formation of the Beaufort Group in South Africa

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.

<span class="mw-page-title-main">Anteosaurinae</span> Extinct subfamily of therapsids

Anteosaurinae is an extinct subfamily of dinocephalian therapsids. It is one of two subfamilies in the family Anteosauridae, the other being Syodontinae.

<i>Sinophoneus</i> Extinct genus of therapsids

Sinophoneus is an extinct genus of carnivorous dinocephalian therapsid belonging to the family Anteosauridae. It lived 272 to 270 million years ago at the beginning of the Middle Permian in what is now the Gansu Province in northern China. It is known by a skull of an adult individual, as well as by many skulls of juvenile specimens. The latter were first considered as belonging to a different animal, named Stenocybus, before being reinterpreted as immature Sinophoneus. Sinophoneus shows a combination of characters present in other anteosaurs. Its bulbous profile snout and external nostrils located in front of the canine are reminiscent of the basal anteosaur Archaeosyodon, while its massive transverse pterygoids processes with enlarged distal ends are more similar to the more derived anteosaurs Anteosaurus and Titanophoneus. First phylogenetic analyzes identified Sinophoneus as the most basal Anteosaurinae. A more recent analysis positioned it outside the Anteosaurinae and Syodontinae subclades, and recovers it as the most basal Anteosauridae.

<i>Bulbasaurus</i> Extinct genus of dicynodonts

Bulbasaurus is an extinct genus of dicynodont that is known from the Lopingian epoch of the Late Permian period of what is now South Africa, containing the type and only species B. phylloxyron. It was formerly considered as belonging to Tropidostoma; however, due to numerous differences from Tropidostoma in terms of skull morphology and size, it has been reclassified the earliest known member of the family Geikiidae, and the only member of the group known from the Tropidostoma Assemblage Zone. Within the Geikiidae, it has been placed close to Aulacephalodon, although a more basal position is not implausible.

Vetusodon is an extinct genus of cynodonts belonging to the clade Epicynodontia. It contains one species, Vetusodon elikhulu, which is known from four specimens found in the Late Permian Daptocephalus Assemblage Zone of South Africa. With a skull length of about 18 centimetres (7.1 in), Vetusodon is the largest known cynodont from the Permian. Through convergent evolution, it possessed several unusual features reminiscent of the contemporary therocephalian Moschorhinus, including broad, robust jaws, large incisors and canines, and small, single-cusped postcanine teeth.

References

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See also