Tchoiria Skeleton
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''Tchoiria'' ('')'' is a
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
neochoristodera Neochoristodera is a lineage of specialised crocodile-like fully aquatic choristodere reptiles. Noted for their long jaws and large size, these animals were predominant across the Northern Hemisphere, occurring in freshwater and coastal environme ...
n
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
of
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
. The name ''Tchoiria'' comes from the city of
Choir A choir ( ), also known as a chorale or chorus (from Latin ''chorus'', meaning 'a dance in a circle') is a musical ensemble of singers. Choral music, in turn, is the music written specifically for such an ensemble to perform or in other words ...
which is nearby to where the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
was found. ''Tchoiria'' is thought to have a similar diet to another neochoristoderan reptile, ''
Champsosaurus ''Champsosaurus'' is an extinct genus of crocodile-like choristodere reptile, known from the Late Cretaceous and early Paleogene periods of North America and Europe (Campanian–Paleocene). The name ''Champsosaurus'' is thought to come from , () ...
'', due to morphology of the skull. It would hunt in freshwater environments, like the living
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family (biology), family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males ...
s, where it would prey on many different types of fish and turtles.


History of research

''Tchoiria'' remains were first recovered as a part of the Joint Soviet-Mongolian Expeditions which took place in the
Gobi Desert The Gobi Desert (, , ; ) is a large, cold desert and grassland region in North China and southern Mongolia. It is the sixth-largest desert in the world. The name of the desert comes from the Mongolian word ''gobi'', used to refer to all of th ...
. They were described by Mikhail B. Efimov in 1975; basing his description on a partial cranium and some parts of the postaxial skeleton found at the Hühteeg Formation. He would make the type species ''T. namsari''. Efimov would name two other ''Tchoiria'' species in the latter 20th century, ''T. magnus'' in 1979 and ''T. egloni'' in 1983. Both taxa were based on postcranial fossils that were also found in the Hühteeg Formation. These would later be redescribed as new members of Neochoristodera by Efimov. The second valid species of ''Tchoiria'' would be named in 2005 by Danial T. Ksepka. ''T. klauseni'' would be based on a partial skull and some postcranial material found at the Two Volcanoes locality of southern Mongolia. Efimov would place ''Tchoiria'' in the order
Choristodera Choristodera (from the Greek χωριστός ''chōristos'' + δέρη ''dérē'', 'separated neck') is an extinct order of semiaquatic diapsid reptiles that ranged from the Middle Jurassic, or possibly Triassic, to the Miocene (168 to 20 or p ...
in his original description. Later it would also be placed in suborder Neochoristodera in 2007 by Ryoko Matsumoto. Matsumoto would declare the suborder
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
in his paper on the choristodere ''
Monjurosuchus ''Monjurosuchus'' is a genus of choristoderan reptile that lived in what is now China and Japan during the Early Cretaceous. It has large eyes, a rounded skull, robust legs with short claws, and a long, thin tail. Fossils have been found that pr ...
''. ''Tchoiria'' was the first choristodere known from Asia and has been used in many morphological and environmental studies based on the order since its original description. There has been a total of four species of ''Tchoiria'' named, but as of 2023, only two are considered valid. ''T. magnus'' was moved to the genus ''Ikechosaurus'' in 1981 and ''T. egloni'' was moved to the new genus '' Irenosaurus'' in 1988. These two taxa were moved due to morphological differences when compared to the ''T. namsari''.


Description


Skull

The skull of ''Tchoiria'' is similar to other neochoristoderes in having a flared postorbital region, an elongated and narrow snout, and having small orbits. The skull is also very flattened like other members of Neochoristodera. The nasals have also fused into a single element. The teeth of ''Tchoiria'' are conical in shape and decrease in size further back in the skull. They also curve medially and have vertical, parallel lines. The orbits have a raised rim surrounding them and are very rough in texture. Like other diapsid reptiles, ''Tchoiria'' has two temporal fenestrae behind the orbit. The snout is shorter than the ''Champsosaurus'' but still longer than other choristoderes. Other skull traits include a shortened lower jaw symphysis, a broader and shorter rostrum, and the posterior displacement of mandibular articulation. ''T. namsari'' is the type species and is known from the Hühteeg Formation of Mongolia. The other species, ''T. klauseni'' is known from the Two Volcanoes locality of southern Mongolia. ''T. klauseni'' was separated from ''T. namsari'' by the number of teeth in both the maxilla and the dentary. ''T. namsari'' is known to have more teeth, 60 in the maxilla and 17 in the symphyseal portion of the dentary, while ''T. klausnei'' has a smaller number of them, 34 in the maxilla and 12 in the symphyseal portion of the dentary.


Postcranial skeleton

A complete set of vertebrae for ''Tchoiria'' have not been fully preserved but specimens of each section have been described. The vertebrae have amphiplatyan-style centrum and a closed notochordal canal. The dorsal vertebrae have small processes below their respective postzygapophyses. These are similar to the processes seen in both '' Simeodosaurus'' and ''
Ikechosaurus ''Ikechosaurus'' is an extinct genus of choristodere reptile which existed in China and Mongolia during the Early Cretaceous. It contains the species ''Ikechosaurus sunailinae'' and ''Ikechosaurus gaoi''. It belongs to the crocodilian-like clade ...
.'' The caudal vertebrae have the unique trait of having a deep groove bordered by ventral flanges; a trait seen in choristoderes but not usually in neochoristoderes. The caudal vertebrae also have unfused hemal arches. ''Tchoiria'' had similar
gastralia Gastralia (: gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these reptil ...
to other members of its suborber. The limbs of ''Tchoiria'' are very fragmentary. Parts of the shoulder girdle are preserved and shows that they would be long and narrow. Besides a single femur, the rest of the hindlimbs are unknown.


Classification

''Tchoiria'' has been placed in Neochoristodera, which is a clade within Choristodera. The group first appeared in the Early Cretaceous of Asia and were successful due to the absence of
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
s. Later in the Cretaceous and the early
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
, neochoristoderans needed to become more specialized because of their shared ecosystem with a wide range of crocodiles. The phylogenetic tree below comes from Dong et al. (2020) shows Neochoristodera and their relationship to other members of Choristodera.


Paleoenvironment

The Khuren Dukh Formation has been sampled as containing mostly
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from ''shale'' by its lack of fissility.Blatt, H., and R.J. Tracy, 1996, ''Petrology.'' New York, New York, ...
layers. This does not change till the upper parts of the formation, transitioning to
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
and
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone and shale. Most of the particles of which the stone is composed are less than and are too small to ...
layers. This formation was first explored in the early 1970's as a part of the Joint Soviet-Mongolian expeditions and has been dated to
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
in age by pollen and invertebrate fossils. The scant plant fossils found nearby shows the environment to be a staple of the Cretaceous; a temperate and plant-covered ecosystem. ''Tchoiria'' lived in a diverse ecosystem of other invertebrate and vertebrate taxa. The aquatic environment included but is not limited to
mollusks Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The num ...
,
ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...
s,
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
,
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s, some crocodiles, and ''Tchoiria'' itself. The terrestrial environment matched many other in the Early Cretaceous. Many dinosaurs including ''
Psittacosaurus ''Psittacosaurus'' ( ; "parrot lizard") is a genus of extinct ceratopsian dinosaur from the Early Cretaceous of what is now Asia, existing between 125 and 105 million years ago. It is notable for being the most species-rich non-avian dinosaur ...
'', ''
Harpymimus ''Harpymimus'' is a basal ornithomimosaurian theropod dinosaur from the Early Cretaceous Period (geology), Period of what is now Mongolia. Unlike later, more derived ornithomimosaurs, ''Harpymimus'' still possessed teeth, although they appear to ...
'', and some unidentified
theropods Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
. They coexisted with some
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s and small
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s. It would prey on fish and turtles mostly and would be hunted by a few crocodiles and theropod dinosaurs. ''Tchoiria'' would be a relatively uncommon animal in its ecosystem, but this could be due to a
sampling bias In statistics, sampling bias is a bias (statistics), bias in which a sample is collected in such a way that some members of the intended statistical population, population have a lower or higher sampling probability than others. It results in a b ...
with the locality.


References

{{Taxonbar, from=Q587956 Aquatic reptiles Choristodera Early Cretaceous reptiles of Asia Paleontology in Mongolia Prehistoric reptile genera