Nochnitsa

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Nochnitsa
Temporal range: Permian, 265–252  Ma
Nochnitsa geminidens holotype.png
Holotype block, containing skull and partial skeleton
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Clade: Gorgonopsia
Genus: Nochnitsa
Kammerer and Masyutin, 2018
Type species
Nochnitsa geminidens
Kammerer and Masyutin, 2018

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.

Contents

Phylogenetic analyzes published since its official description consider it as the most basal gorgonopsian known, due to several anatomical characteristics wo are not present in more or less derived genera. The Vanyushonki Member, the exact site from which Nochnitsa was discovered, would have been a moist, well-vegetated landscape, which would have been periodically flooded. The site contains numerous taxa of contemporary tetrapods, including other various therapsids. The presence of large therocephalians and the smaller size of Nochnitsa and its close relative Viatkogorgon indicate that the latter occupied comparatively small predatory roles.

Discovery and naming

Map showing the Kotelnich locality in Russia where Nochnitsa was found (lower middle left box, magnified in inset box at upper right) Kotelnich locality.jpg
Map showing the Kotelnich locality in Russia where Nochnitsa was found (lower middle left box, magnified in inset box at upper right)

The only known specimen of Nochnitsa, cataloged KPM 310, was discovered in 1994 by the Russian paleontologist Albert J. Khlyupin in the Red Beds of Kotelnich, located along the Vyatka River in Kirov Oblast, European Russia. This specimen was found more precisely in the Vanyushonki Member, a site already known for the discovery of other contemporary therapsids, including the gorgonopsian Viatkogorgon . The datation of this site is not clear, but it seems to date to the latest Guadalupian or early Lopingian epochs. After this discovery, the specimen was subsequently prepared in the Paleontological Museum of Vyatka  [ ru ] by Olga Masyutina. [1]

In 2018, paleontologists Christian F. Kammerer and Vladimir Masyutin named new genera of gorgonopsians and therocephalians discovered at Kotelnitch in two articles in the scientific journal PeerJ . [1] [2] In their paper focusing on gorgonopsians, the specimen KPM 310 is identified as the holotype of a new genus and species, which they name Nochnitsa geminidens. [1]

Nochnitsa is named after the Nocnitsa, a nocturnal hag-like creature from Slavic mythology. Its name was intended as a parallel to the Gorgons, similarly hag-like creatures from Greek mythology, which are the namesake of many genera within Gorgonopsia and the clade as a whole. The name also reflects the nocturnal habits inferred for the genus. The type species name, geminidens, means "twin tooth" and refers to one of the autapomorphies of the species, postcanine teeth arranged in pairs. [1]

Description

Skull

Nochnitsa is small for a gorgonopsian, with a skull only 82 millimetres (3.2 in) long. It had a relatively long snout with five incisors, a canine, and six postcanine teeth on each side. The postcanine teeth are autapomorphic for the genus in being arranged in three pairs of closely placed teeth separated by longer diastemata. In each pair, the posterior tooth is larger. The mandible is relatively slender and lacks a strong "chin", unlike other gorgonopsians. [1]

Postcranial skeleton

Right side of the block containing the holotype specimen, showing in more detail the rare postcranial elements known of the animal Nochnitsa geminidens holotype right side.png
Right side of the block containing the holotype specimen, showing in more detail the rare postcranial elements known of the animal

Although incompletely known, the holotype specimen of Nochnitsa contains part of the postcranial elements with the skull, including the cervical vertebrae, some dorsal vertebrae, and associated ribs. The right forelimb is also preserved and partially articulated. [1]

In the cervical vertebrae, the axial spine is broadly rounded and similar in morphology to that of other gorgonopsians. The dorsal vertebrae are preserved as central and transverse process fragments interspersed by the ribs. The ribs are also simple and elongated. The scapula is elongated, narrow and weakly curved, comparable to that of other gorgonopsians of similar size like Cyonosaurus , but different from the anteroposteriorly broadened scapular spines of Inostrancevia . [1]

The humerus is relatively slender, having a short, poorly developed delto-pectoral ridge, where the muscles attach to the upper arm. The radius and ulna, have a distinct distal curvature, and the distal tip of the radius forms a discrete differentiated rim of the shaft. No olecranon process is visible on the ulna, but it is possible that this is the result of a lesion. [lower-alpha 1] The preserved proximal carpal elements consist of the radial, the ulnar and two smaller, irregular elements that would probably represent the centralia. The ulnar is the longest carpus on the proximodistal side and is widened at its proximal and distal ends. The radial is a shorter and more rounded element. The possible centralia, although poorly preserved, appear to be weakly curved. The concave surface of the centralia would presumably have been articulated with the radial, based on the conditions of other gorgonopsians. [lower-alpha 2] Several small irregular bones between the proximal carpals and the metacarpals probably represent distal carpals, but these elements are too poorly preserved to be further identified. Based on their great length relative to the other manual elements, the two best preserved elements probably represent the third and fourth metacarpals, which are the longest of all other gorgonopsians for which the manus are known. A shorter but still elongated element may represent the fifth metacarpal. A semi-articulated set of poorly preserved bones appear to represent fingers, one potentially ending in the ungual. Based on the size of the phalanx-like elements, these probably correspond to the third and fourth fingers, disarticulated from the third and fourth metacarpals. These elements are too poor for a definitive count of the phalanges, and there is no clear evidence of the reduced disc-shaped phalanges commonly present in gorgonopsians. [1]

Classification

Skulls of Viatkogorgon and Nochnitsa (reversed for comparison) Viatkogorgon and Nochnitsa skulls.png
Skulls of Viatkogorgon and Nochnitsa (reversed for comparison)

Nochnitsa is currently the most basal gorgonopsian known, and its position is justified by several plesiomorphic criteria, such as the lowered mandibular symphysis, the low and inclined front of the dentary bone (similar to those of therocephalians), as well as a surface and a row of elongated teeth. These mentioned features are not present in derived genera. [1] The 2018 analysis by Kammerer and Masyutin, although derived from a previous analysis conducted by one of the two authors, [3] is a major revision of the phylogeny of the gorgonopsians, discovering that the derived representatives are divided into two groups, of Russian and African origin. [1] The basal position of Nochnitsa in phylogenetic analysis of gorgonopsians is still recognized in later published studies. [4] [5]

The following cladogram showing the position of Nochnitsa within Gorgonopsia follows Kammerer and Rubidge (2022): [5]

Gorgonopsia

Nochnitsa

Viatkogorgon

"Russian clade"

Suchogorgon

Sauroctonus

Pravoslavlevia

Inostrancevia

"African clade"

Phorcys

Eriphostoma

Gorgonops

Cynariops

Lycaenops

Smilesaurus

Arctops

Arctognathus

Rubidgeinae

Paleoecology

Paleoenvironment

Life restoration of N. geminidens Nochnitsa restoration.jpg
Life restoration of N. geminidens

Nochnitsa is known from the Kotelnich locality, which consists of a series of Permian red bed exposures along the banks of the Vyatka River in Russia. It is specifically from the Vanyushonki Member, which is the oldest rock unit in the Kotelnich succession, consisting of pale or brown mudstones (clay and silts, with some fine-grained sand) as well as gray mudstone, and dark red mudstone at the base of this exposure. These mudstones were possibly deposited from suspension in standing water bodies on floodplains or shallow ephemeral lakes, that remained flooded for short periods of time, but the exact environment has not yet been determined, due to the lack of a primary structure of the sediments. The presence of rootlets, roots and tree stumps would show that the landscape represented by the member of Vanyushonki would be relatively humid and well vegetated. Although the age of the Kotelnitch faunal complex is uncertain, it may date to the same age as those found in South Africa, which date from either the Late Middle Permian or the Early Late Permian. [1] [6]

The Vanyushonki Member contains abundant fossils of tetrapods contemporary to Nochnitsa, most including numerous fossils often consisting of articulated and complete skeletons. Apart from its close relative Viatkogorgon, other therapsids from the locality include the anomodont Suminia and the therocephalians Chlynovia , Gorynychus , Karenites , Perplexisaurus , Scalopodon , Scalopodontes , and Viatkosuchus . The pareiasaur Deltavjatia is particularly abundant there, and the parareptile Emeroleter is present. [1] [2] [7] Fossil ostracods have also been found. [6]

Ecological niche

As the fossil record shows, the fauna of Kotelnitch was mainly dominated by the large therocephalians, and more specifically by Gorynychus and Viatkosuchus. These two taxa being much larger than Nochnitsa and Viatkogorgon, this indicates that the gorgonopsians occupied smaller predatory roles than the large therocephalians. This is further confirmed by the fact that several gorgonopsians having appeared after the extinction of the end of the Guadalupian reach considerably larger sizes than the two previously mentioned genera. [2] [4] This type of ecological niche is also similar to that seen in the Pristerognathus Assemblage Zone in the Karoo Basin, South Africa, prior to the main round of gorgonopsian diversification there. [2] However, he noted that some Guadalupian gorgonopsians, notably Phorcys , are already larger in size, indicating that not all genera shared similar roles. [5]

See also

Notes

  1. The proximal end of this feature is not complete and was partially replaced by mudstone during the fossilization of the holotype specimen. [1]
  2. A clear intermediate is not visible, as this element is generally small in gorgonopsians and may be absent or still buried in the fossil block containing the holotype specimen. [1]

Related Research Articles

<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.

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

Inostrancevia is an extinct genus of large carnivorous therapsids who 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.

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

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.

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

Trochosaurus is a dubious genus of therocephalian therapsid from South Africa based upon specimens of Lycosuchidae. Three species of Trochosaurus have appeared in literature, T. intermedius, T. major, and T. dirus. The genus Trochosaurus and all three species are considered nomen dubia today, while the specimens referred to them are regarded as Lycosuchidae incertae sedis.

Angelosaurus is an extinct genus of herbivorous caseid synapsids that lived during the late Lower Permian (Kungurian) and early Middle Permian (Roadian) in what is now Texas and Oklahoma in the United States. Like other herbivorous caseids, it had a small head, large barrel-shaped body, long tail, and massive limbs. Angelosaurus differs from other caseids by the extreme massiveness of its bones, particularly those of the limbs, which show a strong development of ridges, processes, and rugosities for the attachment of muscles and tendons. Relative to its body size, the limbs of Angelosaurus were shorter and wider than those of other caseids. The ungual phalanges looked more like hooves than claws. The few known cranial elements show that the skull was short and more robust than that of the other representatives of the group. Angelosaurus is also distinguished by its bulbous teeth with shorter and wider crowns than those of other caseids. Their morphology and the high rate of wear they exhibit suggests a diet quite different from that of other large herbivorous caseids, and must have been based on particularly tough plants. A study published in 2022 suggests that the genus may be paraphyletic, with Angelosaurus possibly only represented by its type species A. dolani.

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

Sauroctonus is an extinct genus of gorgonopsian therapsids who lived during the end of the Middle Permian in what is now European Russia. The first fossils, discovered in Tatarstan, were first considered to belong to a new species of the South African genus Arctognathus, named A. progressus in 1938. The taxon will be designated as such until 1940, when it will be assigned to the genus Inostrancevia by Ivan Yefremov, before being definitively classified in a separate genus erected by Alexey Bystrow in 1955. The most complete known fossils of S. progressus include cranial and postcranial elements currently all recorded in Tatarstan. These elements show that the animal is a rather medium-sized gorgonopsian.

<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>Rubidgea</i> Extinct genus of therapsids

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.

<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>Pravoslavlevia</i> Extinct genus of therapsids

Pravoslavlevia is an extinct genus of gorgonopsian therapsids that lived in the late Permian and is part of the Sokolki subcomplex of Russia. It had a skull 22 centimetres (8.7 in) long. The total length of the animal was about 1.4 metres. Only one species is known.

<i>Megawhaitsia</i> Extinct genus of large therapsids from the Late Permian in East Russia

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.

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

Eriphostoma is an extinct genus of gorgonopsian therapsids known from the Middle Permian of Tapinocephalus Assemblage Zone, South Africa. It has one known species, Eriphostoma microdon, and was first named by Robert Broom in 1911. It is the oldest known gorgonopsian and among the smallest and most basal members of the clade.

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

Smilesaurus is an extinct genus of gorgonopsian known from Africa. It lived during the Late Permian. It contains the single species S. ferox.

"Dixeya" nasuta is a species of the predatory gorgonopsian therapsids from the Late Permian of East Africa, known from fossils found in what is now Tanzania. The species has a complicated taxonomic history, it was originally named as a second species of the genus Dixeya which is now considered a junior synonym of Aelurognathus. "D." nasuta itself, however, was not moved to Aelurognathus, and although it was instead tentatively referred to Arctognathus at first it has since been recognised to not belong to this genus either. This situation leaves "Dixeya" nasuta without a formal genus name. It was proposed to belong to a new distinct genus, named "Njalila", that was informally proposed for the species in a PhD thesis, but this name has not yet been formally published and is currently a nomen nudum. "D." nasuta has been characterised from other gorgonopsians by a combination of its straight snout profile, upturned and 'pinched' nose, and curved jaw margin.

<i>Karenites</i> Genus of therapsids from the Late Permian of Russia

Karenites is an extinct genus of therocephalian therapsids from the Late Permian of Russia. The only species is Karenites ornamentatus, named in 1995. Several fossil specimens are known from the town of Kotelnich in Kirov Oblast.

<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.

<i>Gorynychus</i> Genus of therapsids from the mid-Permian of Russia

Gorynychus is a genus of therocephalian from the mid-Permian from Kotelnich, Russia. The genus contains two species, G. masyutinae and G. sundyrensis. It was named after the three-headed dragon Zmey Gorynych from Russian mythology.

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.

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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  2. 1 2 3 4 Christian F. Kammerer; Vladimir Masyutin (2018). "A new therocephalian (Gorynychus masyutinae gen. et sp. nov.) from the Permian Kotelnich locality, Kirov Region, Russia". PeerJ . 6: e4933. doi: 10.7717/peerj.4933 . PMC   5995100 . PMID   29900076.
  3. Christian F. Kammerer (2016). "Systematics of the Rubidgeinae (Therapsida: Gorgonopsia)". PeerJ . 4: e1608. doi: 10.7717/peerj.1608 . PMC   4730894 . PMID   26823998.
  4. 1 2 Eva-Maria Bendel; Christian F. Kammerer; Nikolay Kardjilov; Vincent Fernandez; Jörg Fröbisch (2018). "Cranial anatomy of the gorgonopsian Cynariops robustus based on CT-reconstruction". PLOS ONE . 13 (11): e0207367. doi: 10.1371/journal.pone.0207367 . PMC   6261584 . PMID   30485338.
  5. 1 2 3 Christian F. Kammerer; Bruce S. Rubidge (2022). "The earliest gorgonopsians from the Karoo Basin of South Africa". Journal of African Earth Sciences . 194: 104631. Bibcode:2022JAfES.19404631K. doi:10.1016/j.jafrearsci.2022.104631. S2CID   249977414.
  6. 1 2 Michael J. Benton; Andrew J. Newell; Al'bert Y. Khlyupin; Il'ya S. Shumov; Gregory D. Price; Andrey A. Kurkin (2012). "Preservation of exceptional vertebrate assemblages in Middle Permian fluviolacustrine mudstones of Kotel'nich, Russia: stratigraphy, sedimentology, and taphonomy". Palaeogeography, Palaeoclimatology, Palaeoecology . 319–320: 58–83. Bibcode:2012PPP...319...58B. doi:10.1016/j.palaeo.2012.01.005.
  7. Elena G. Kordikova; Albert J. Khlyupin (2001). "First evidence of a neonate dentition in pareiasaurs from the Upper Permian of Russia". Acta Palaeontologica Polonica . 46 (4): 589–594. Archived from the original on 27 January 2022. Retrieved 27 January 2022.