Leninia
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''Leninia'' 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 basal
ophthalmosaurine Ophthalmosaurinae is an extinct subfamily of ophthalmosaurid thunnosaur ichthyosaurs from the Middle Jurassic to the late Early Cretaceous (Bajocian - Albian) of Europe, North America and South America. Currently, the oldest and the basalmost kno ...
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
known from the late
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
(lower
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
stage) of western
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
. ''Leninia'' was first named by Valentin Fischer, Maxim S. Arkhangelsky, Gleb N. Uspensky, Ilya M. Stenshin and
Pascal Godefroit Pascal Godefroit is a Belgian paleontologist. He discovered dinosaurs like ''Olorotitan'' in 2003. Godefroit is the director of earth and life sciences at the Royal Belgian Institute of Natural Sciences The Museum of Natural Sciences of Belgium ...
in
2013 File:2013 Events Collage V2.png, From left, clockwise: Edward Snowden becomes internationally famous for leaking classified NSA wiretapping information; Typhoon Haiyan kills over 6,000 in the Philippines and Southeast Asia; The Dhaka garment fact ...
and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
is ''Leninia stellans''. It was named for
Vladimir Lenin Vladimir Ilyich Ulyanov. ( 1870 – 21 January 1924), better known as Vladimir Lenin,. was a Russian revolutionary, politician, and political theorist. He served as the first and founding head of government of Soviet Russia from 1917 to 19 ...
, one of the leaders of the
Communist Revolution A communist revolution is a proletarian revolution often, but not necessarily, inspired by the ideas of Marxism that aims to replace capitalism with communism. Depending on the type of government, socialism can be used as an intermediate stage ...
in Russia, as the museum it is currently housed in is also named for him. The specimen was discovered in 2012, on the banks of the river
Volga 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 ...
near the town of Kriushi in a limestone nodule. There are several other fossils from the same time period near it, including
ammonites Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...
, lamellibranchs and fish remains. Only part of the skull was found.


Features

The
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
is incomplete, and without any
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
. it is partially crushed transversely, and nothing further forward than the
naris A nostril (or naris , plural ''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 turbi ...
remains, with what seems to be a fairly clean break. It is not known whether this was due to
orogeny Orogeny is a mountain building process. An orogeny is an event that takes place at a convergent plate margin when plate motion compresses the margin. An ''orogenic belt'' or ''orogen'' develops as the compressed plate crumples and is uplifted t ...
and rock action or to the ichthyosaur's death. The specimen is around long, which suggests the skull was between long overall. Unusually, the posterior process of the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
extends as far back as the
orbit In celestial mechanics, an orbit 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 object or position in space such as a p ...
, and about halfway along it, possibly even projecting into the orbit at points. The maxillary dental groove is extremely shallow, only deep. The lacrimal-
nasal Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination: * With reference to the human nose: ** Nasal administration, a method of pharmaceutical drug delivery ** ...
contact is elongated, so that the lacrimal, rather than the prefrontal, forms the posterior edge of the naris. The anterior process and shaft of the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
are unusually narrow, but the posterior plate is well-developed and extends as high as the middle of the orbit. The
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
is small compared to the orbit, with a large dorsal orbital rim extending out from the top of the bone. The
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
seems more robust than that of other ophthalmosaurines, but has a thin anterior surface which articulates with the postorbital. There is a concave area which a ligament would connect to the quadrate process. The
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 The quadratojugal is a skull bone present in many vertebrates, including ...
is present but incomplete. However, it is possible to tell the shape by the marks where it would have articulated with other bones. Above the naris, the nasal bone forms a conspicuous lateral 'wing'. There is no clear evidence for a
foramen In anatomy and osteology, a foramen (;Entry "foramen"
in
near the prefrontal-nasal facet. The prefrontal forms a thick anterior process which has grown over the nasal, unlike in other ophthalmosaurids. Its suture with the lacrimal bone is straight. The frontals are roughly triangular and have the 'internasal' foramen between them (not between the nasals). They have long posterior processes which suture with the forked processes of the parietals in a star shape, hence the specific name ''stellans''. The
postfrontal The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
contacts the nasal over a short distance, and as in ''Ophthalmosaurus,'' has no Y-shaped anterior process. Uniquely, the postfrontal does not border the
supratemporal fenestra The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, but still forms a long process in contact with the supratemporal. The parietals touch the anteriomedial process of the
supratemporal The supratemporal bone is a paired cranial bone present in many tetrapods and tetrapodomorph fish. It is part of the temporal region (the portion of the skull roof behind the eyes), usually lying medial (inwards) relative to the squamosal and latera ...
s, another
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
. This anteromedial process also blocks the postfrontal from bordering the supratemporal fenestra. The parietals are crushed against one another, but there is no evidence of a crest. The supratemporal is more extensive than in most ophthalmosaurines, but is separated from the postorbital by the postfrontal and squamosal bones. Both pterygoids are present but cannot really be seen, and are distorted. The
basioccipital The basilar part of the occipital bone (also basioccipital) extends forward and upward from the foramen magnum, and presents in front an area more or less quadrilateral in outline. In the young skull this area is rough and uneven, and is joined t ...
is not perfectly preserved, but the extracondylar area is reduced and concave. The
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
is bulbous, but the floor of the
foramen magnum The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblon ...
is not well preserved enough to be a diagnostic feature. The occipital head of the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foot ...
is largely expanded, with a large hyoid process, but the quadrate head is missing. The quadrate is roughly ear-shaped, with an occipital lamella. Only the posterior end of the
lower jaw 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 ...
is preserved, and this lacks any distinctive features, such as teeth. The surangular fossa is present but extremely reduced, and the angular is well exposed laterally. These features, however, are not diagnostic or autapomorphies. Both
sclerotic ring Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
s are preserved, but the left one is much less distorted. It is composed of 14 trapezoidal plates, with
crenulation In a geological context, crenulation or crenulation cleavage is a fabric formed in metamorphic rocks such as phyllite, schist and some gneiss by two or more stress directions causing the formation of the superimposed foliations. Formation Crenula ...
s on the internal edge. The sclerotic aperture is only 11.03% of the orbital area, suggesting that the specimen was fully grown. (All of this section is citing )


Evolutionary importance

''Leninia'' is one of the latest-living ophthalmosaurines, and also one of the most basal. Despite this, it still had the very large sclerotic ring and aperture, suggesting that this was one of the most basal characteristics of ophthalmosaurines and that they were all deep-diving, remaining conservative in this
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
during the entire time period that they existed. The absolute size of ''Leninia's'' sclerotic ring is among the largest known in ichthyosaurs, with only ''
Baptanodon ''Baptanodon'' is an ichthyosaur of the Late Jurassic period (160-156 million years ago), named for its supposed lack of teeth (although teeth of this genus have since been discovered). It had a graceful long dolphin-shaped body, and its jaws we ...
'' ('''Ophthalmosaurus natans')'' and the giant ''
Temnodontosaurus ''Temnodontosaurus'' (Greek for "cutting-tooth lizard"temno, meaning "to cut", odont meaning "tooth" and sauros meaning "lizard") is an extinct genus of ichthyosaur from the Early Jurassic period. They lived between 200 and 175 million years ag ...
'' exceeding it. As all the ophthalmosaurines had this kind of structure, and were more similar in eye structure than the platypterygines, this suggests that they did not diversify in the same way that the platypterygines did but stayed similar as deep divers, possibly as they were so specialised for deep diving that they were outcompeted in other niches.


Phylogeny

Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
based in the phylogenetic analysis of Fischer ''et al.'', 2013: The following cladogram shows a possible phylogenetic position of ''Leninia'' in Ophthalmosauridae according to the analysis performed by Zverkov and Jacobs (2020).


See also

*
List of ichthyosaurs This list of ichthyosaurs is a comprehensive listing of all genera that have ever been included in the order Ichthyosauria or the parent clade Ichthyopterygia, excluding purely vernacular terms. The list includes all commonly accepted genera, but a ...
*
Timeline of ichthyosaur research This timeline of ichthyosaur research is a chronological listing of events in the history of paleontology focused on the ichthyosauromorphs, a group of secondarily aquatic marine reptiles whose later members superficially resembled dolphins, shar ...
*
List of organisms named after famous people (born 1800–1899) In biological nomenclature, organisms often receive scientific names that honor a person. A taxon (e.g. species or genus; plural: taxa) named in honor of another entity is an eponym, eponymous taxon, and names specifically honoring a person or pers ...


References

{{Taxonbar, from=Q6336273 Early Cretaceous ichthyosaurs Fossil taxa described in 2013 Extinct animals of Russia Vladimir Lenin Ophthalmosaurinae Ichthyosauromorph genera