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''Suminia'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of basal
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. Anomodonts were very diverse during the Middle Pe ...
that lived during the Tatarian age of the late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
, spanning approximately from 268–252 Ma.Rybczynski N. 2000. Cranial anatomy and phylogenetic position of Suminia getmanovi, a basal anomodont (Amniota: Therapsida) from the Late Permian of Eastern Europe. Zool. J. Linn. Soc. 130:329–73 ''Suminia'' is recognized as the youngest non-
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
anomodontIvachnenko MF. 1994. A new Late Permian dromasaurian (Anomodontia) from Eastern Europe. Paleontological Journal 28: 96- 103. and was named in honor of Russian paleontologist D. L. Sumin. Its fossil localities are primarily derived from the Kotel’nich locality of the
Kirov Oblast Kirov Oblast ( rus, Кировская область, p=ˈkʲirəfskəjə ˈobləsʲtʲ) is a federal subject of Russia (an oblast) located in Eastern Europe. Its administrative center is the city of Kirov. As of the 2010 census, the population ...
in
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
. However, there have been some isolated specimens found in a few different localities, all from eastern European regions of Russia.Fröbisch, J. and Reisz, R. R. 2011. The postcranial anatomy of Suminia getmanovi (Synapsida: Anomodontia), the earliest known arboreal tetrapod. Zoological Journal of the Linnean Society, 162: 661–698. ''Suminia'', along with '' Otsheria'' and '' Ulemica'' make up the monophyletic group of Russian basal anomodonts named
Venyukovioidea Venyukovioidea is an infraorder of anomodont therapsids related to dicynodonts from the Permian of Russia. They have also been known as Venjukovioidea, as well as by the similar names Venyukoviamorpha or Venjukoviamorpha in literature. This in pa ...
.Modesto, S. & B. Rubidge (2000) A basal anomodont therapsid from the lower Beaufort Group, Upper Permian of South Africa, Journal of Vertebrate Paleontology, 20:3, 515-521. These Venyukovioid anomodonts are understood to have been derived from an ancestor that dispersed from Gondwana into Euramerica.Modesto, S. P., B. S. Rubidge, and J. Welman. 1999. The most basal anomodont therapsid and the primacy of Gondwana in the evolution of the anomodonts. Proc. R. Soc. Lond. B 266:331–337. ''Suminia getmanovi'' is the only defined species within the genus and it is known for specializations in teeth for effective, functional oral processing of plant material as well as being one of the first species with a proposed
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
lifestyle.Fröbisch, J. and Reisz, R. R. 2009. The Late Permian herbivore Suminia and the early evolution of arboreality in terrestrial vertebrate ecosystems. Proceedings of the Royal Society B, 276: 3611–3618.


Discovery

''Suminia'' was named and described by Russian paleontologist M. F. Ivakhnenko in 1994, based on a nearly complete skeleton (PIN 2212/10) discovered by Russian paleontologist S. N. Getmanov on a dig led by D. L. Sumin in 1990 on the bank of the Vyatka River near the town of Kotel'nich in Kirov Oblast, Russia. Skulls, lower jaws, and isolated teeth found in the same region were also referred to the genus.Ivachnenko MF. 1994. A new Late Permian dromasaurian (Anomodontia) from Eastern Europe. Paleontological Journal 28: 96- 103. New material of ''Suminia'' was excavated from the Upper Permian Kotel’nich locality in which a single large block with articulated skeletons of at least 15 specimen of ''Suminia getmanovi'' was found. This new material of ''Suminia getmanovi'' consisted of mainly subadult to adult specimens that were well preserved (no signs of weathering or predation), suggesting rapid burial perhaps due to a catastrophic event.


Description

''Suminia'' are diagnosed as a small Venyukoviid with various autapomorphies such as large orbit (almost 1/3 of the skull length), large teeth in relation to the skull size, reduced number of 23 presacral vertebrae, a more long than wide cervical vertebrae (suggests elongated neck), elongated limbs, manus and pes equal to ~40% of length of their respective limb, and enlarged distal carpal 1 and tarsal 1.


Cranial anatomy

''Suminia getmanovi'' are recognized for their well-preserved skulls and teeth. ''Suminia'' skull length is fairly small, measuring in at 58mm long characterized with a short snout with its
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...
regions expanded. While the
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
composes of around 27% of the total skull length, the external
naris A nostril (or naris , : nares ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbinates, ...
is also large, measured to compose of about 13% of the total skull length. Cranial features that are only shared with ''Ulemica'' that distinguish ''Suminia'' and ''Ulemica'' from other anomodonts is the preparietal absence, a reduced interparietal suture located anterior to the pineal foramen, and narrow palatine. ''Suminia'' cranial anatomy can also be defined by their raised pineal foramen (in comparison with other taxa with the pineal foramen flush with the skull) and premaxilla contact with palatine, which are all features shared by its infraorder, Venyukovioidea. Perhaps one of the most striking cranial anatomy features of Suminia is its similarity in masticatory architecture with dicynodonts, indicating that the sliding jaw articulation may have originated before dicynodonts. ''Suminia'' dentition has significant implications on its feeding ecology, which is discussed below.


Postcranial anatomy

Study of ''Suminia'' postcranial anatomy reveals many autapomorphies for the single species. Significant postcranial autapomorphies of ''Suminia'' are the reduced number of presacral and dorsal vertebrae (exclusively amphicoelous) with lack of fusion in the sacral region between vertebrae (suggests high flexibility), wide pre- and postzygapophyses, longer proportions of cervical pleurocentra, distinct proportionally longer limbs, a manus that forms ~ 40% of the length of the forelimb with particularly long, curved terminal phalanges, a pes that makes up ~38% of the hindlimb, and enlarged carpal 1 and tarsal 1 (suggests divergent first digit). These different morphological features indicate a significantly deviated postcranial anatomy from other anomodonts, suggesting that ''Suminia'' adopted an arboreal lifestyle (see below).


Feeding ecology

In combination with a masticatory architecture similar to ''Dicynodonts'' (defined by sliding jaw articulation) ''Suminia''’s canineless, large leaf-shaped teeth follow occluding dentition that is completely marginal, which differs from other species with leaf-shaped teeth present. This provides indications not only for
herbivory A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
, but into the mechanisms of oral processing.Ivakhnenko, M.F. Cranial morphology and evolution of Permian Dinomorpha (Eotherapsida) of eastern Europe. Paleontol. J. 42, 859–995 (2008). One interesting feature of the occlusal pattern in ''Suminia'' dentition is the angle of the occluding surfaces. With an angle of 75 degrees from jaw plane, it is suggested that ''Suminia''’s more posterior teeth shred food material rather than crush it. The anterior teeth are observed to be significantly larger and devoid of this occlusal pattern. Therefore, the more anterior teeth are suggested to be responsible for cutting off pieces of plant for the posterior teeth to shred. ''Suminia'' are therefore understood to have been obligatory herbivores as the dentition and mandibular function permits shredding of plant material via posterior translation. Teeth replacement was discovered to be infrequent, which was shown in certain specimens of ''Suminia getmanovi'' that were seen to have teeth worn down almost to the neck. However, these wear facets on the upper and lower posterior teeth are in themselves, consistent with herbivory in that the locations of the wear (labial and lingual aspects of upper and lower posterior teeth respectively), are distinct evidence that the wear isn't a consequence of tooth-to-food but rather tooth-to-tooth occlusion.Rybczynski, N. & Reisz, R. R. 2001 Earliest evidence for efficient oral processing in a terrestrial herbivore. Nature 411, 684 –687. The evidence for ''Suminia''’s extensive oral processing suggest that ''Suminia'' dentition is highly specialized for high fiber herbivory. This provides an alternative explanation that the ability to process tough, high fiber plant material may have been a more basal feature of anomodonts than previously thought.


Habitat/lifestyle

Adaptations to arboreal lifestyle are understood to evolve through convergent evolution. However, many arboreal vertebrates share similar physical mechanisms (grasping, clinging, hooking).Hildebrand, M. & Goslow, G. E. J. 2001 Analysis of vertebrate structure, 5th edn. New York, NY: John Wiley & Sons. Other features common to climbing and tree-living animals include lighter bones, longer limbs, small size, and a longer tail. The
varanopids Varanopidae is an extinct family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards (with the name of the group deriving from the monitor lizard genus ''Varanus'') and may have filled a similar niche. T ...
'' Eoscansor'' from the Pennsylvanian subperiod of the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
of North America and ''
Ascendonanus ''Ascendonanus'' (meaning "climbing dwarf") is an extinct genus of varanopid amniote from the Early Permian of Germany. It is one of the earliest specialized arboreal (tree-living) tetrapods currently known and outwardly resembled a small lizar ...
'' from the Lower
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
of Germany show earlier anatomical specializations for climbing and life in trees among synapsids. ''Suminia'' shares many features with other arboreal
tetrapods A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
. While its forelimbs retain the sprawling posture found in other anomodonts, the digits of both front and back paws were enlarged and modified for climbing. Evidence for grasping abilities include a first digit that could spread away from the others. The first digit is measured to have an angle of ~30-40 degrees to the remaining digits of the manus and pes, which grants ability of the first digit to flex ventrally, independent of the rest of the digits. This is further supported by the elongated limbs and claw shaped, laterally compressed terminal phalanges which would aid in clinging ability. In addition, the tail anatomy with expansion of the anterior region and suggests ability of balance as well as
prehensile Prehensility is the quality of an appendage or organ that has adapted for grasping or holding. The word is derived from the Latin term ''prehendere'', meaning "to grasp". The ability to grasp is likely derived from a number of different origin ...
, grasping abilities, providing more evidence for arboreal lifestyle. A study of the microanatomy of ''Suminia''′s limb bones published in 2025 added more support for a tree-living lifestyle. Research revealed relatively thin bone walls and a lack of internal support structures called
trabeculae A trabecula (: trabeculae, from Latin for 'small beam') is a small, often microscopic, tissue element in the form of a small beam, strut or rod that supports or anchors a framework of parts within a body or organ. A trabecula generally has a ...
, in strong contrast to the very thick bone walls and dense trabeculae typically found in other anomodonts. Along with its small size, ''Suminia''′s lighter limb bones would be an adaptation for climbing in trees. Via a morphometric analysis as well as comparison to other arboreal vertebrates, ''Suminia getmanovi'' provides anatomical evidence that it lived among the trees, stamping a significant mark in evolutionary history for arboreal lifestyle.


Classification

''Suminia'' belong to the monophyly/infraorder Venyukovioidea, a sub clade of basal anomodonts.


See also

*
List of therapsids This list of therapsids is an attempt to create a comprehensive listing of all Genus, genera that have ever been included in the Therapsida excluding mammals and purely vernacular terms. The list includes all commonly accepted genera, but also gene ...


References

{{Taxonbar, from=Q135229 Anomodont genera Lopingian synapsids of Europe Prehistoric synapsids of Europe Fossil taxa described in 1994 Fossils of Russia