Megawhaitsia

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Megawhaitsia
Temporal range: Wuchiapingian
Megawhaitsia maxilla.jpg
Illustration of the holotype maxilla showing possible venom ducts
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Clade: Therocephalia
Family: Whaitsiidae
Genus: Megawhaitsia
Ivakhnenko, 2008
Type species
Megawhaitsia patrichae

Megawhaitsia is an extinct genus of large therocephalian therapsids who lived during the Late Permian (Wuchiapingian) in what is now Eastern Europe. The only known species is M. patrichae, described in 2008 from several fossils discovered in various oblasts of European Russia. The fossils are representative of a large animal whose skull size is estimated to be 40–50 cm (16–20 in) long.

Contents

The most notable feature of Megawhaitsia is that it has a maxilla with canals directly connected to the tooth root of the canines. Based on the characteristics present in the related genus Euchambersia , Russian paleontologist Mikhail Ivakhnenko raises the possibility that the animal may have had a venom gland. If it is true, then it would then be one of the oldest tetrapods known to have this attribute. Subsequent studies have challenged this proposition.

The imposing size of Megawhaitsia and its position as an apex predator could be linked to the disappearance or absence of large gorgonopsians at the end of the Late Permian in certain regions of present-day European Russia. Megawhaitsia could thus have occupied the ecological niches previously occupied by the gorgonopsians.

Discovery and naming

The holotype specimen of Megawhaitsia was discovered in the mid-1950s during excavations carried out in the locality of Vyazniki-2, located in Vladimir Oblast, in European Russia, before being cataloged as PIN 1100/101. This site is dated to the Wuchiapingian stage of the Late Permian. [1] It had originally been incorrectly recorded as the jawbone of a gorgonopsian similar to Inostrancevia . Given the low presence of gorgonopsians during the Late Permian in Russia, the fossil was reassigned to a large therocephalian in a work published in 1997, without however receiving a binomial name. [2] In 2001, Mikhail Feodosievich Ivakhnenko attributed two additional fossil remains to the still unnamed taxon. The first is a partial maxillary bone, cataloged PIN 1538/39, discovered in the locality of Purly, in the Nizhny Novgorod Oblast. The second is the right part of an incomplete mandible, cataloged PIN 4417/101, discovered in the locality of Shabarshata, in the Kirov Oblast. [3]

Ivakhnenko published in 2008 a formal description of the new taxa, based on the specimens PIN 1100/101 and PIN 1538/39, describing it as the first Late Permian whaitsiid from Eastern Europe. [1] Subsequently, the scope of whaitsiids was expanded to include discoveries made earlier in the Permian deposits of the same Russian regions, including Moschowhaitsia and Viatkosuchus , described in 1963 and 1995 respectively. [4] The genus name Megawhaitsia comes from the Ancient Greek μέγας (megas, "great"), combined with the name of another therocephalian genus, Whaitsia (name witch is today synonymous with Theriognathus ). The specific epithet patrichae honors the Australian paleontologist Patricia Vickers-Rich. [1]

Description

Restoration of the head Moschorhinid1DB.jpg
Restoration of the head

The known fossils of Megawhaitsia are very incomplete, thus preventing any complete reconstruction of the animal's anatomy. However, the structure of the two known maxillary bones proves that it is a therocephalian. [2] [3] [1]

The animals' maxillary bone was massive with the largest preserved fragment measuring about 10 cm (3.9 in) in length. Based on the proportions of the fuller skulls of the smaller South African representatives of the group, the total length of the animal's skull is estimated to be between 40–50 cm (16–20 in), which would make Megawhaitsia the largest therocephalian known to date. On the lower edge of the maxilla there is a large space that can accommodate the lower canine. Behind are the sockets of three large upper canines, the two anterior being somewhat larger. The roots of the teeth are deep, all three having a deep alveolar fossa. [1] The partial mandible attributed to Megawhaitsia has the socket of a very large canine but lacks those of the cheek teeth. [3]

A feature of the maxillary bone is that it has three channels which start in the region of the lacrimo-nasal duct, pass along the roots of the teeth and open near the sockets of each of the canines. By analogy with the hypotheses on the venomousness of another genus of therocephalians, Euchambersia , Ivakhnenko interprets these canals as a possible proof of the presence of poisonous glands in Megawhaitsia, which would be used to slaughter large prey. [1] However, since the venomousness of Euchambersia has been questioned in a study published in 2017, in particular on the basis of the comparison with various modern venomous animals, the authors of the 2017 study suggest other explanations of the presence of these maxillary canals might be possible. [5]

Classification

During the second half of the 20th century, the fossil maxillary bones of Megawhaitsia were considered to belong to a gorgonopsid similar or identical to the genus Inostrancevia. In 1997, the fossils were reassigned to an undetermined therocephalian in the family Whaitsiidae, [2] then to the Moschorhinidae family in 2001. [3] In the 2008 article of Ivakhnenko, Megawhaitsia is included again in the family Whaitsiidae, within the superfamily Whaitsioidea. At that time, the Whaitsioidea taxon included the Euchambersiidae and the Whaitsiidae as sister-groups, due to their similar appearance. [1] A study published less than a year later by Adam Huttenlocker estimated that the families Euchambersiidae, Moschorhinidae and Annatherapsididae represented junior synonyms of Akidnognathidae, considered the sister-group of Whaitsiidae. [6] It was in 2016 that Huttenlocker and Christian Sidor concluded that the Akidnognathidae are in fact close to the Chthonosauridae, the two forming the sister-group of a clade containing the Whaitsioidea and the Baurioidea. [7] The superfamily Whaitsioidea remains recognized as a valid taxa, although it now only contains whaitsiids and a few related genera. [4] [7]

Paleobiology

In comparison to South African therocephalians, Megawhaitsia had a noticeably larger size corresponding to a specialized carnivorous predator niche. It fed on fairly large prey, notably dicynodonts, which were numerous in Russian regions of Europe during the Upper Permian. The possible presence of venom glands in Megawhaitsia would be consistent with the warm-blooded dicynodont hypothesis, as venom offers a significant advantage especially in hunting active warm-blooded prey. [1]

One of the types of large coprolites found in the Vyazniki locality is associated with Megawhaitsia or closely related whaitsiids such as Moschowhaitsia. It reveals a high content of bony material, including bones bearing traces of a rich network of blood vessels, probably belonging to dicynodonts, indicating a predator that occupied the top position in the trophic chain. Additionally, remains of fish scales and material interpreted as ganoine have been found in morphotype A coprolites, as well as fur-like structures. These are interpreted as the oldest fossil coat remains known to date, although it remains unclear whether they belong to prey or were swallowed by a predator as a result of grooming. [8]

Paleoecology

The locality of Viazniki-2, where the holotype of Megawhaitsia was discovered, contains numerous fossils of tetrapods dating from the Wuchiapingian, including the temnospondyl Dvinosaurus , as well as non-amniote reptiliomorphs, including the seymouriamorph Karpinskiosaurus and numerous chroniosuchians. Sauropsids present include pareiasaurs such as Obirkovia and archosauriforms of the family Proterosuchidae, such as Archosaurus . The latter would also have been one of the main predators of the area. Other therapsids are present in the locality, such as an indeterminate dicynodont and even other therocephalians, including Annatherapsidus , Malasaurus and Moschowhaitsia. [3] [1]

Researchers speculate that due to their increased size, East European whaitsiids occupied the ecological niche of the large gorgonopsians, which at that time had disappeared from Eastern Europe, possibly due to a climate cooling. [1]

See also

Related Research Articles

<span class="mw-page-title-main">Therapsid</span> Clade of tetrapods including mammals

Therapsida is a major group of eupelycosaurian synapsids that includes mammals, 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.

<span class="mw-page-title-main">Gorgonopsia</span> Extinct group of saber-toothed therapsids from the Permian

Gorgonopsia is an extinct clade of sabre-toothed therapsids from the Middle to Upper Permian roughly 265 to 252 million years ago. They are characterised by a long and narrow skull, as well as elongated upper and sometimes lower canine teeth and incisors which were likely used as slashing and stabbing weapons. Postcanine teeth are generally reduced or absent. For hunting large prey, they possibly used a bite-and-retreat tactic, ambushing and taking a debilitating bite out of the target, and following it at a safe distance before its injuries exhausted it, whereupon the gorgonopsian would grapple the animal and deliver a killing bite. They would have had an exorbitant gape, possibly in excess of 90°, without having to unhinge the jaw.

<span class="mw-page-title-main">Biarmosuchia</span> Extinct suborder of therapsids

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.

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

Inostrancevia is an extinct genus of large carnivorous therapsids which lived during the Late Permian in what is now European Russia and South Africa. The first known fossils of this gorgonopsian were discovered in the Northern Dvina, where two almost complete skeletons were exhumed. Subsequently, several other fossil materials were discovered in various oblasts, and these finds will lead to a confusion about the exact number of valid species in the country, before only three of them were officially recognized : I. alexandri, I. latifrons and I. uralensis. More recent research carried out in South Africa has discovered fairly well-preserved remains of the genus, being attributed to the species I. africana. The whole genus is named in honor of Alexander Inostrantsev, professor of Vladimir P. Amalitsky, the paleontologist who described the taxon.

<span class="mw-page-title-main">Therocephalia</span> Extinct order of therapsids

Therocephalia is an extinct clade of eutheriodont therapsids from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their teeth, suggest that they were carnivores. Like other non-mammalian synapsids, therocephalians were once described as "mammal-like reptiles". Therocephalia is the group most closely related to the cynodonts, which gave rise to the mammals. This relationship takes evidence in a variety of skeletal features.

<span class="mw-page-title-main">Theriodontia</span> Clade of therapsids

The theriodonts are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts.

<i>Euchambersia</i> Extinct genus of therapsid from Late Permian South Africa

Euchambersia is an extinct genus of therocephalian therapsids that lived during the Late Permian in what is now South Africa and China. The genus contains two species. The type species E. mirabilis was named by paleontologist Robert Broom in 1931 from a skull missing the lower jaw. A second skull, belonging to a probably immature individual, was later described. In 2022, a second species, E. liuyudongi, was named by Jun Liu and Fernando Abdala from a well-preserved skull. It is a member of the family Akidnognathidae, which historically has also been referred by as the synonymous Euchambersiidae.

<i>Moschorhinus</i> Genus of synapsid from late Permian and early Triassic South Africa

Moschorhinus is an extinct genus of therocephalian in the family Akidnognathidae with only one species: M. kitchingi. It was a carnivorous synapsid which has been found in the Late Permian to Early Triassic of the South African Karoo Supergroup. It was a large carnivore, reaching 1.5 m (4.9 ft) in total body length with the largest skull comparable to that of a lion in size. It had a broad, blunt snout which bore long, straight canines. It appears to have replaced the gorgonopsids ecologically, and hunted much like a big cat. While most abundant in the Late Permian, it survived a little after the Permian Extinction, though these Triassic individuals had stunted growth.

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

Viatkogorgon is a genus of gorgonopsian that lived during the Permian period in what is now Russia. The first fossil was found at the Kotelnich locality near the Vyatka River and was made the holotype of the new genus and species V. ivachnenkoi in 1999. The generic name refers to the river and the related genus Gorgonops—the gorgons of Greek mythology are often referenced in the names of the group. The specific name honors the paleontologist Mikhail F. Ivakhnenko. The holotype skeleton is one of the most complete gorgonopsian specimens known and includes rarely preserved elements such as gastralia and a sclerotic ring. A larger, but poorly preserved specimen has also been assigned to the species.

<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>Scylacops</i> Extinct genus of therapsids

Scylacops is an extinct genus of Gorgonopsia. It was first named by Broom in 1913, and contains two species, S. bigendens, and S. capensis. Its fossils have been found in South Africa and Zambia. It is believed to be closely related to the Gorgonopsian Sauroctonus progressus. Scylacops was a moderately sized Gorgonopsid.

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

Venyukovia is an extinct genus of venyukovioid therapsid, a basal anomodont from the Middle Permian of Russia. The type and sole species, V. prima, is known only by a partial lower jaw with teeth. Venyukovia has often been incorrectly spelt as 'Venjukovia' in English literature. This stems from a spelling error made by Russian palaeontologist Ivan Efremov in 1940, who mistakenly replaced the 'y' with a 'j', which subsequently permeated through therapsid literature before the mistake was caught and corrected. Venyukovia is the namesake for the Venyukovioidea, a group of small Russian basal anomodonts also including the closely related Otsheria, Suminia, Parasuminia and Ulemica, although it itself is also one of the poorest known. Like other venyukovioids, it had large projecting incisor-like teeth at the front and lacked canines, although the remaining teeth are simple compared to some other venyukovioids, but may resemble those of Otsheria.

<i>Moschowhaitsia</i> Extinct genus of therapsid from the late Permian of Eurasia

Moschowhaitsia is an extinct genus of therocephalian therapsids from the Late Permian (Guadalupian) of Russia and China. The type species, Moschowhaitsia vjuschkovi, was discovered in the Changxingian-aged Archosaurus Assemblage Zone of Russia and named in 1963 by Russian palaeontologist Leonid Petrovich Tatarinov. A second species was discovered in Jingtai County of Gansu, China in 2020 and named as M. lidaqingi in 2023 by Jun Liu and Fernando Abdala, the first whaitsiid therocephalian to be discovered in China. It was among the larger carnivores in the faunal assemblages it occurred in, with a skull-length of up to 25 centimetres (9.8 in) in M. vjuschkovi and an even larger estimated 35 centimetres (14 in) for M. lidaqingi, one of the largest therocephalian skulls reported. The genus name Moschowhaitsia alludes to two other therocephalians, Moschorhinus and Whaitsia, due to the structure of its palate combining physical features of both these genera.

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

Akidnognathidae is an extinct family of therocephalian therapsids from the Late Permian and Early Triassic of South Africa, Russia and China. The family includes many large-bodied therocephalians that were probably carnivorous, including Moschorhinus and Olivierosuchus. One akidnognathid, Euchambersia, may even have been venomous. Akidnognathids have robust skulls with a pair of large caniniform teeth in their upper jaws. The family is morphologically intermediate between the more basal therocephalian group Scylacosauridae and the more derived group Baurioidea.

Ichibengops is an extinct genus of therocephalian therapsids known from the type species Ichibengops munyamadziensis, which lived in what is now Zambia during the Late Permian. Ichibengops was named in 2015 on the basis of fossils found in the Wuchiapingian-age Madumabisa Mudstone Formation in the Luangwa Basin. Therocephalians have been known from the Luangwa Basin for decades, yet Ichibengops was the first endemic Zambian therocephalian to have been described in detail. Phylogenetic analysis indicates that it is a basal member of the clade Eutherocephalia, lying just outside a clade containing hofmeyriids, whaitsiids, and baurioids. Ichibengops is the sister taxon of the Russian therocephalian Chthonosaurus; together they form one of several known African-Russian sister taxon pairs of eutherocephalians, which indicate that eutherocephalians could freely disperse across most of Pangea during the Late Permian. Like the fellow therocephalian Euchambersia, Ichibengops might have had venom glands, as evidenced by grooves above its teeth.

<span class="mw-page-title-main">Rubidgeinae</span> Extinct subfamily of mammals

Rubidgeinae is an extinct subfamily of gorgonopsid therapsids known only from Africa. They were among the largest gorgonopsians, and their fossils are common in the Cistecephalus and Daptocephalus assemblage zones of the Karoo Basin. They lived during the Late Permian, and became extinct at the end of the Permian.

Leogorgon is a extinct genus of dubious therapsid from the Late Permian Sokolki Faunal Assemblage of Russia. It was originally classified as a rubidgeine gorgonopsian, and would have been the first member of that clade from outside of Africa if that identification had been valid. However, it may instead be a combination of the tooth of a gorgonopsian and the braincase of a dicynodont, and may be a wastebin taxon.

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

Nochnitsa is an extinct genus of gorgonopsian therapsids who lived during an uncertain stage of the Permian in what is now European Russia. Only one species is known, N. geminidens, described in 2018 from a single specimen including a complete skull and some postcranial remains, discovered in the red beds of Kotelnich, Kirov Oblast. The genus is named in reference to Nocnitsa, a nocturnal creature from Slavic mythology. This name is intended as a parallel to the Gorgons, which are named after many genera among gorgonopsians, as well as for the nocturnal behavior inferred for the animal. The only known specimen of Nochnitsa is one of the smallest gorgonopsians identified to date, with a skull measuring close to 8 cm (3.1 in) in length. The rare postcranial elements indicate that the animal's skeleton should be particularly slender.

Phorcys is an extinct genus of gorgonopsian that lived during the Middle Permian period (Guadalupian) of what is now South Africa. It is known from two specimens, both portions from the back of the skull, that were described and named in 2022 as a new genus and species P. dubei by Christian Kammerer and Bruce Rubidge. The generic name is from Phorcys of Greek mythology, the father of the Gorgons from which the gorgonopsians are named after, and refers to its status as one of the oldest representatives of the group in the fossil record. Phorcys was recovered from the lowest strata of the Tapinocephalus Assemblage Zone (AZ) of the Beaufort Group, making it one of the oldest known gorgonopsians in the fossil record—second only to fragmentary remains of an indeterminate gorgonopsian from the older underlying Eodicynodon Assemblage Zone.

References

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