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''Parasumina'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
anomodont Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: Ra ...
known from the late Capitanian
age Age or AGE may refer to: Time and its effects * Age, the amount of time someone or something has been alive or has existed ** East Asian age reckoning, an Asian system of marking age starting at 1 * Ageing or aging, the process of becoming older ...
at the end of the
middle Permian 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. ...
period of
European Russia European Russia (russian: Европейская Россия, russian: европейская часть России, label=none) is the western and most populated part of Russia. It is geographically situated in Europe, as opposed to the cou ...
. The type and only species is ''Parasuminia ivakhnenkoi''. It was closely related to '' Suminia'', another Russian anomodont, and was named for its resemblance ("similar to ''Suminia''"). Little is known about ''Parasuminia'' as the only fossils are of fragmentary pieces of the skull and jaw, but the known remains suggest that its head and jaws were deeper and more robust than those of ''Suminia'', and with shorter, stouter teeth. However, despite these differences they appear to have been similar animals with a similarly complex method of processing vegetation.


Description

''Parasuminia'' is only known from incomplete remains of the lower jaw and the front of the snout. Compared to ''Suminia'', the lower jaw is deeper and more robust, and is down-turned towards its tip with a deep, rounded 'chin'. The teeth in both the upper and lower jaws are lower and wider than in ''Suminia'', with serrated edges only found on the newest replacement teeth before being worn away. The tooth row consists of 10 to 12 teeth and is straight along its surface, with longer procumbent (forward projecting) incisors at the tip of the lower jaw to accommodate the down-turned tip. Likewise, the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
from the front of the upper jaw is also taller than in ''Suminia'' and with relatively shorter teeth. The tips of the upper jaw are similarly deflected upwards, opposing the down-turned lower jaw, and the front teeth are also procumbent. Although incomplete, the total length of the skull was estimated to be roughly long, slightly larger than the long skull of ''Suminia''. The
parietal bone The parietal bones () are two bones in the Human skull, skull which, when joined at a fibrous joint, form the sides and roof of the Human skull, cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, an ...
from the back of the skull roof is larger and proportionately wider than it is in ''Suminia'', and completely surrounds the circular pineal foramen (or "third eye"). The pineal foramen itself is raised slightly above the surface of the skull; however, it is less raised than the chimney-like structure in ''Suminia'' and has more gently sloping sides. In contrast with the smooth surface of the rest of the parietal, the bone around the pineal foramen has a rugose texture, imprinted with blood vessels.


History of discovery

Fossils of ''Parasuminia'' are known exclusively from a locality known as Sundyr-1 near the mouth of the Sundyr River (a tributary to the
Volga River The Volga (; russian: Во́лга, a=Ru-Волга.ogg, p=ˈvoɫɡə) is the List of rivers of Europe#Rivers of Europe by length, longest river in Europe. Situated in Russia, it flows through Central Russia to Southern Russia and into the Cas ...
) in the
Mari El Republic The Mari El Republic (russian: Респу́блика Мари́й Эл, ''Respublika Mariy El''; Meadow Mari: ; Hill Mari: ) is a republic of Russia. It is in the European Russia region of the country, along the northern bank of the Volga Rive ...
of European Russia, and were first discovered in 2009 by an expedition from the Borissiak Paleontological Institute of the
Russian Academy of Sciences The Russian Academy of Sciences (RAS; russian: Росси́йская акаде́мия нау́к (РАН) ''Rossíyskaya akadémiya naúk'') consists of the national academy of Russia; a network of scientific research institutes from across t ...
,
Moscow Moscow ( , US chiefly ; rus, links=no, Москва, r=Moskva, p=mɐskˈva, a=Москва.ogg) is the capital and largest city of Russia. The city stands on the Moskva River in Central Russia, with a population estimated at 13.0 million ...
. The site itself was originally discovered by a group of schoolboys in 1997, and the site and its fossils were briefly reported on by A. Yu. Bezerin in 2005 who referred to the site as the "Yul’yaly locality". The site was later renamed Sundyr-1 and interpreted to belong to a new stratigraphic
member Member may refer to: * Military jury, referred to as "Members" in military jargon * Element (mathematics), an object that belongs to a mathematical set * In object-oriented programming, a member of a class ** Field (computer science), entries in ...
, the Ustpoldarsa Member, of the Poldarsa Formation. The fossil fauna recovered from Sundyr-1 have been collated into the Sundyr Assemblage Zone, interpreted as representing an intermediate fauna between the older Middle Permian ' dinocephalian faunas' and later ' theriodontian faunas' of the Late Permian. The Sundyr Assemblage has been interpreted as belonging to the lower Upper Severodvinian stage in Russian stratigraphy, correlating to the upper Capitanian stage of the International Stratigraphic Scale dating to approximately 259 million years ago. The remains of ''Parasuminia'' were initially referred to as ' aff. ''Suminia, and were later described as a distinct genus and
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
in 2017 by A. A. Kurkin. The
holotype specimen A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, PIN no. 5388/196, consists of a partial pair of lower jaws with teeth. Other specimens include numerous lower jaw fragments, a single partial premaxilla and many isolated teeth. An isolated right parietal, PIN no. 5388/198, may also belong to ''Parasuminia''. The remains of ''Parasuminia'' currently consist only of fragmented, disarticulated pieces of skull as well as isolated pairs of
dentaries In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
that remain held together by strong suturing at the jaw tips. The generic name is from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
"''para''" ("close", "similar") and the genus ''Suminia'' for its close resemblance and relationship to the latter. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
is in memory of the
palaeontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
M.F. Ivakhnenko and in recognition of his "outstanding" work on Russian palaeontology.


Classification

''Parasuminia'' was assigned to the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Galeopidae by Kurkin, a family originally erected solely for the
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
n anomodont ''
Galeops ''Galeops'' is an extinct genus of anomodont therapsids from the Middle-Late Permian of South Africa. It was described by Robert Broom in 1912. Some cladistic analyses have recovered it as closely related to dicynodonts. See also * List of ther ...
'' by palaeontologist
Robert Broom Robert Broom FRS FRSE (30 November 1866 6 April 1951) was a British- South African doctor and palaeontologist. He qualified as a medical practitioner in 1895 and received his DSc in 1905 from the University of Glasgow. From 1903 to 1910, he ...
in 1912. ''Galeops'' was formerly included in a group of small anomodonts known as the '
dromasaurs Dromasaurs are a paraphyletic group of anomodont therapsids from the Middle Permian. They were small with slender legs and long tails. Their skulls were short, but the eye sockets were large. Dromasauria was once considered to be a major group ...
' together with '' Galechirus'' and ''
Galepus ''Galepus'' is an extinct genus of anomodont therapsids. See also * List of therapsids This list of therapsids is an attempt to create a comprehensive listing of all genera that have ever been included in the Therapsida excluding mammals ...
'', however, phylogenetic analyses of anomodonts have since shown that 'dromasaurs' are polyphyletic and so are not a natural group of related species. However, Kurkin assigned ''Parasuminia'' to Galeopidae based on an alternative
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
classification for anomodonts (as well as other synapsids) proposed by Russian palaeontologist M. F. Ivakhnenko. Under Ivakhnenko's taxonomy, the family Galeopidae consists of the traditional 'dromasaurs' as well as other anomodonts, including ''Suminia'' and '' Anomocephalus'', and is classed within the
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Dromasaurida, itself under the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typicall ...
ia proper. Ivakhnenko proposed galeopids were grouped together for their use a palinal jaw stroke (pulling their lower jaw backwards to chew), a trait they share with dicynodonts which he considered to indicate they shared a common ancestry. This opposes modern orthodox phylogenetic classifications of anomodonts, which do not group these anomodonts together within a single clade. ''Parasuminia'' has yet to be included in a phylogenetic analysis of anomodonts, and so its evolutionary relationships outside of this scheme are unclear. However, ''Parasuminia'' was referred to the Venyukovioidea, a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
that includes ''Suminia'' but not any of the 'dromasaurs', by palaeontologists Kenneth Angielczyk and Christian Kammerer in 2018.


Palaeobiology

Like other related anomodonts, ''Parasuminia'' would have been a herbivore. Due to the similarity of the wear marks left on the teeth of ''Parasuminia'' to those of ''Suminia'', it is likely that they both fed in a similar fashion with extensive tooth-on-tooth occlusion. Furthermore, although the jaw joint of ''Parasuminia'' is unknown, it can be inferred from the pattern of wear that ''Parasuminia'' was capable of a palinal jaw stroke to shred vegetation. This complex method of chewing is shared with both ''Suminia'' and the dicynodonts.


Palaeoecology

In the Sundyr Assemblage fauna, ''Parasuminia'' coexisted with two large carnivorous
therocephalian Therocephalia is an extinct suborder of eutheriodont therapsids (mammals and their close relatives) from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their te ...
s, the predatory scylacosaurid '' Julognathus'' and the suggested scavenger '' Gorynychus sundyrensis''. Large herbivores are poorly known, and are only represented by dinocephalians possibly similar to '' Ulemosaurus''. A number of early tetrapods are also known from this assemblage, including ''
Leptoropha ''Leptoropha'' is an extinct genus of aquatic seymouriamorph known from the Middle Permian of Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the larges ...
'' aff. ''talonophora'', '' Microphon'', '' Enosuchus'' sp., and the
chroniosuchia Chroniosuchia is a group of tetrapods that lived from the Middle Permian to Late Triassic in what is now Eastern Europe, Kyrgyzstan, China and Germany. Chroniosuchians are often thought to be reptiliomorphs, but some recent phylogenetic analyses ...
n ''
Suchonica ''Suchonica'' is an extinct genus of chroniosuchid reptiliomorph from upper Permian (upper Tatarian age) deposits of Sukhona Formation of Vologda Region, Russia. It was first named by V. K. Golubev in 1999 File:1999 Events Collage.png, From ...
'', as well as the
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
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 ...
''
Dvinosaurus ''Dvinosaurus'' is an extinct genus of amphibious temnospondyls localized to regions of western and central Russia during the middle and late Permian, approximately 265-254 million years ago. Its discovery was first noted in 1921 by Russian pal ...
'' sp. The Sundyr Assemblage fauna has been considered to be a 'crisis fauna', representing the last of the dinocephalian dominated faunas in Eurasia at the end of the Middle Permian and transitioning into the theriodontian dominated faunas of the Late Permian.


References

{{Taxonbar, from=Q39050744 Anomodont genera Guadalupian synapsids of Europe Fossils of Russia Fossil taxa described in 2017