Konzhukovia
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''Konzhukovia'' is an
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
genus that belongs to an extinct group of
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
, the largest clade of basal
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s including about 198 genera, 292 species, and more than half of which were alive during the early
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
period. The animal was a predator that lived about 260 million years ago, and could get up to about three meters in length. Specifically, ''Konzukovia'' lived during the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
, between 252 and 270 million years ago according to the type of rock the fossil was found in. There are three species within this genus, ''K. vetusta'', ''K. tarda'', and ''K. sangabrielensis'', the first two originating from Russia while the latest originating from Southern Brazil. The discovery of this specimen in Southern Brazil provided more evidence to support the idea that during this animals existence, there was a “biological corridor” because of the supercontinent
Pangea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
, allowing these species to be found so far apart from each other. ''Konzhukovia'' belongs to the family
Archegosauridae Archegosauridae is a family of relatively large and long snouted temnospondyls that lived in the Permian period. They were fully aquatic animals, and were metabolically and physiologically more similar to fish than modern amphibians.Florian Witzm ...
, a family consisted of large temnospondyls that most likely compare to modern day crocodiles. Since the discovery of the latest species, ''K. sangabrielensis'', Pacheco proposes that there must be the creation of a new family, Konzhokoviidae, a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group in a sister-group relationship with Stereospondlyi in order to accommodate the three species. ''Konzhukovia'' skulls usually exhibit typical rhinesuchid features including an overall parabolic shape, small orbits located more posteriorly, and the pterygoids do not reach the
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...
. These animals were long-snouted amphibians that had clear adaptations made for fish catching, as well as exemplifying aquatic features.


Etymology

The name ''Konzhukovia,'' was given because of its “rhinesuchid-like” temnospondyl properties. It was assigned the name ''Konzohuvia'' by Schoch and Milner. The specific name ''sangabrielensis'' (a species type) is derived from the municipality where the specimen was recovered in southern Brazil.


Paleobiology

The ''Konzhukovia'' skull has an overall triangular appearance due to the lateral margins of the snout expanding parallel and then diverging from each other and the most anterior portion of the snout is rounded. The skull contains both infra- and supraorbital sensory sulci, the infraorbital sulcus is located on the maxilla and runs posteriorly to the lacrimal while the supraorbital sulcus extends anteriorly to the naris. The orbits are elongate and the skull shows paired anterior palatal vacuity. The orbits are positioned after the midline of the skull and are relatively widely separated from each other. The skull contains pleurodont dentition and the maxilla and premaxilla of recovered specimens carry evidence of more than 30 teeth. The tooth row runs straight between the vomerine fangs, the vomerine tusks and palatine tusks are equal in size, and the maxillary teeth are anteroposteriorly compressed. Comparing the three species, ''K. sangabrielensis'' was almost twice the size of both ''K. tarda'' and ''K. vetusta'' and appeared to be very robust in overall shape. Pacheco used phylogenetic inference to infer that ''K. sangabrielensis'' was missing the septomaxilla due to the fact that the absence of this bone is a common feature in its relatives, as it was not visible in the specimen found representing ''K. sangabrielensis''. All three ''Konzhukovia'' species exhibit oval nostrils with wide lateral openings located on the anterior edge of the snout (Pacheco, 2016). The premaxilla, maxilla, and nasal all make contact with the nostrils edges and the prefrontal is fragmented. The maxilla is what forms most of the lateral portion of the choanae, giving an overall long and oval form which is seen in all konzhukoviids and several other temnospondyls. The vomer contacts the maxilla and its lateral process reaches the same level as the palatine tucks. The cultriform process of the parasphenoid in ''K. vetusta and K. tarda'' is inserted between the vomers. These animals have interpterygoid vacuities which appear to be oval and elongated, the palatine is what forms the most anterior part of the vacuity. The pterygoids are fragmented and in ''K. tarda,'' the palatine ramus of the pterygoid runs anteriorly to the palatine tusks, unlike in ''K. vetusta'' where the palatine ramus ends at the same level. The illiac blade is known to be not bifurcated in ''K. vetusta''.


Discovery


''Konzhukovia vetusta''

''Konzhukovia vetusta'' was named in 1955 by E. D. Konzhukova as a ''
Melosaurus ''Melosaurus'' is an extinct genus of temnospondyl amphibian. Fossils of it have been found in Russia. Fossil remains of ''Melosaurus uralensis'' were discovered at Belebei in Russia. Its fossils dated of the Asselian-Olenekian stages (299–24 ...
vetusta''. A ''K. vetusta'' skull was found in the Bolshekinelskaya Formation at the Malyi Uran locality, Orenburg region, in Russia. The ''K. vetusta'' skull was originally described by Gubin, in 1991, and assigned to the family of Melosauridae which later becomes renamed.


''Konzhukovia tarda''

''Konzhukovia tarda'' was found in the
Ocher Assemblage Zone Ochre ( ; , ), or ocher in American English, is a natural clay earth pigment, a mixture of ferric oxide and varying amounts of clay and sand. It ranges in colour from yellow to deep orange or brown. It is also the name of the colours produced ...
in the Orenburg region of Russia.


''Konzhukovia sangabrielensis''

In 2016, Cristian Pereira Pacheco described ''K. sangabrielensis'' from the anterior half and partial right side of a skull roof and palate of a temnospondyl from South America. After diagnosis of the species by Pacheco and his group, it was clear that it was of the genus ''Konzhukovia'' which was previously recorded exclusively from Russia. The specimen of ''K. sangabrielensis'' that is described by Pachecho was recovered from Posto Queimado (Early
Guadalupian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
),
Rio do Rasto Formation The Rio do Rasto Formation is a Late Permian sedimentary geological formation in the South Region of Brazil. The official name is Rio do Rasto, although in some publications it appears as ''Rio do Rastro''. Geography It is found mainly in the B ...
, state of
Rio Grande do Sul Rio Grande do Sul (, , ; "Great River of the South") is a Federative units of Brazil, state in the South Region, Brazil, southern region of Brazil. It is the Federative_units_of_Brazil#List, fifth-most-populous state and the List of Brazilian st ...
, southern Bazil. Initially, this specimen was recorded as a Melosaurinae by Dias-da-Silva (2012), but phylogenetic analysis supported its placement within Tryphosuchinae, basal to the Russian temnospondyls of this group. Once ''K. sangabrielensis'' was discovered, the new family of Konzhokoviidae was proposed in order to accommodate the new Brazilian species in relation to its Russian relatives. ''K. sangabrielensis'' remains the basal most konzhukoviid in the family.


Geological setting

The
Rio do Rasto Formation The Rio do Rasto Formation is a Late Permian sedimentary geological formation in the South Region of Brazil. The official name is Rio do Rasto, although in some publications it appears as ''Rio do Rastro''. Geography It is found mainly in the B ...
, where ''K. sangabrielensis'' was recovered, ranges from Guadalupian to Lopingian. The Permian specimens that have been found in southern South America only consist of fossil faunas in the Rio do Rasto Formation. The Southern Urals area of European Russia, where ''K. tarda'' and ''K. vetusta'' were found, have been the recovery site of many amphibians and reptiles from the Upper Permian, dating back to the 1940s. The area with rich deposits consists of 900,000 km2 of land between River Volga in the northwest and Orenburg in the southwest. A continental succession consisting of mudstones, siltstones, and sandstones provides specimens from the last two stages of the Permian, the Kazanian and Tatarian.


References

{{Taxonbar, from=Q6430467 Permian temnospondyls Prehistoric amphibian genera Fossils of Russia Permian temnospondyls of South America Permian Brazil Fossils of Brazil Paraná Basin Fossil taxa described in 1991