Parasuminia
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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 living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
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 In the geologic timescale, the Capitanian is an age or stage of the Permian. It is also the uppermost or latest of three subdivisions of the Guadalupian Epoch or Series. The Capitanian lasted between and million years ago. It was preceded by ...
age 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 ± ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
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 ''Suminia'' is an extinct genus of basal anomodont that lived during the Tatarian age of the late Permian, spanning approximately from 268-252 Ma.Rybczynski N. 2000. Cranial anatomy and phylogenetic position of Suminia getmanovi, a basal anomodon ...
'', 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 Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
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 ...
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 skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is n ...
from the back of the skull roof is larger and proportionately wider than it is in ''Suminia'', and completely surrounds the circular
pineal foramen A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhyth ...
(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 longest river in Europe. Situated in Russia, it flows through Central Russia to Southern Russia and into the Caspian Sea. The Volga has a length of , and a catch ...
) 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 Dinocephalians (terrible heads) 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. Dinocephal ...
faunas' and later '
theriodontia The theriodonts (clade Theriodontia) 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. Naming In 1876, Ric ...
n faunas' of the
Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
. The Sundyr Assemblage has been interpreted as belonging to the lower Upper Severodvinian
stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...
in Russian stratigraphy, correlating to the upper
Capitanian In the geologic timescale, the Capitanian is an age or stage of the Permian. It is also the uppermost or latest of three subdivisions of the Guadalupian Epoch or Series. The Capitanian lasted between and million years ago. It was preceded by ...
stage of the International Stratigraphic Scale dating to approximately 259 million years ago. The remains of ''Parasuminia'' were initially referred to as '
aff. ''Species affinis'' (commonly abbreviated to: ''sp.'', ''aff.'', or ''affin.'') is taxonomic terminology in zoology and botany. In open nomenclature it indicates that available material or evidence suggests that the proposed species is related to, h ...
''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 appropriat ...
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 sever ...
, 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 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 pe ...
"''para''" ("close", "similar") and the genus ''Suminia'' for its close resemblance and relationship to the latter. The specific name 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 foss ...
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 group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Galeopidae ''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 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 coun ...
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 o ...
' together with ''
Galechirus ''Galechirus'' is an extinct genus of anomodont therapsids. It was about 30 cm (1 ft) long. Description ''Galechirus'' was lizard-like in appearance. It is considered to be a dicynodont by some paleontologists; others think ''Galech ...
'' 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 In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of anomodonts have since shown that 'dromasaurs' are
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
and so are not a natural group of related species. However, Kurkin assigned ''Parasuminia'' to Galeopidae based on an alternative taxonomic classification for anomodonts (as well as other
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s) 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 ''Anomocephalus'' is an extinct genus of primitive anomodonts and belongs to the clade Anomocephaloidea. The name is said to be derived from the Greek word ''anomos'' meaning lawless and ''cephalos'' meaning head. The proper word for head in Gre ...
'', 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 ...
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, typic ...
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 Venyukovioidea is an infraorder of anomodont therapsids related to dicynodonts from the Permian of Russia. They have also known as Venjukovioidea, as well as by the similar names Venyukoviamorpha or Venjukoviamorpha in literature. This in part ...
, 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 ter ...
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 t ...
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 ''Ulemosaurus'' is an extinct genus of dinocephalian therapsids that lived 265 to 260 million years ago, at Isheevo in Russian Tatarstan. It was a tapinocephalid, a group of bulky herbivores which flourished in the Middle Permian. ''Ulemosaurus'' ...
''. A number of early
tetrapods 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 therapsi ...
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 largest c ...
'' aff. ''talonophora'', '' Microphon'', ''
Enosuchus ''Enosuchus'' is an extinct genus of seymouriamorphs from Russia during 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. I ...
'' sp., and the chroniosuchian ''
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, F ...
'', as well as the
temnospondyl 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 ...
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
''
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 pa ...
'' 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