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''Coeruleodraco'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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
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 ...
n known from the Late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
( Oxfordian)
Tiaojishan Formation The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian-Oxfordian (stage), Oxfordian stages). It is known for its Lagerstätte, exceptionally preser ...
in China. ''Coeruleodraco'' is significant as the most complete Jurassic choristodere taxon, as the only other named Jurassic choristodere '' Cteniogenys'' is based on fragmentary remains. Although similar to '' Philydrosaurus'' in its proportions and postcranial characters, it is distinct in retaining several apparently
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...
characters, including a short snout, paired external nares and an open lower temporal fenestra.


Description

The skeleton was small, about 20.3 cm (8.0 inches) in length, not including the tail. The tail is incomplete, but including its estimated length, the total length of the skeleton would have been closer to 40.0 cm (15.7 inches). ''Coeruleodraco'' had a generalized, lizard-like body type. This is similar to some early choristoderes (like '' Cteniogenys'', '' Philydrosaurus'', and '' Monjurosuchus''), but contrasts with other choristodere body types including long-necked forms such as '' Hyphalosaurus'', and
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 ...
-like forms such as ''
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 , () ...
'' and other neochoristoderes. The skull had a short, pointed snout with about 38 teeth on each side (a low number by choristodere standards). It also had two nares (nostril holes). While paired nares are standard in most reptile groups, almost all other choristoderes (except ''Lazarussuchus'') typically have one single hole in the skull for the nostrils, making ''Coeruleodraco'''s retention of paired nares a
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...
("primitive") feature relative to most other choristoderes. In common with other choristoderes, ''Coeruleodraco'' has elongated prefrontal bones which meet each other at the top of the snout, separating a pair of small, triangular nasal bones in the process. The nasals are so reduced in size (especially compared to ''Philydrosaurus'') that they are completely surrounded by the prefrontals and premaxillary bones, and fail to contact the toothed maxillary bones. 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 ...
and
postfrontal The postfrontal is a paired cranial bone found in many tetrapods. It occupies an area of the skull roof between and behind the orbits (eye sockets), lateral to the frontal and parietal bones, and anterior to the postorbital bone. The postfrontal ...
bones (which lie behind the eye) are separate from each other; in many other choristoderes these bones fuse into a single postorbitofrontal bone. The rear of the skull is not particularly unusual, with each side of the skull having two holes known as temporal fenestrae (as with other diapsids). The upper temporal fenestra is not elongated (unlike neochoristoderes), but the lower temporal fenestra is low and long. The rear edge of each upper temporal fenestra was formed by the
squamosal bone 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 ancestral ...
, which was ornamented with small spines. The postcranial skeleton was very similar to that of ''Philydrosaurus''. The neck was short, and its vertebrae had low, swollen neural arches (upper portions). There were six or seven vertebrae in the neck, 16 in the torso, three in the hip, and more than 24 in the tail, which is incomplete. Vertebrae were longer than high, and the neural arches were completely fused to the centra (lower, spool-shaped portions), in contrast to the condition in neochoristoderes. Vertebrae at the base of the tail were fused to fan-shaped ribs which projected straight out. Further down the tail, these ribs became shorter and rectangular. The humerus was robust, with a roughly-textured knob above the large entepicondyle. The plate-like
ischium The ischium (; : is ...
bone (which formed the rear lower part of the hip) had a distinct triangular prong along its rear edge.


Classification

A group of
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
recovered ''Coeruleodraco'' as one of the earliest diverging members of a group of small-bodied Eurasian choristoderes. This group was likely a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
according to the parsimony analysis, in which case it was the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of the gharial-like neochoristoderes, although its
monophyly 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 comm ...
is more uncertain under the bootstrap analysis. The only known genus of choristodere which diverged earlier than ''Coeruleodraco'' was '' Cteniogenys'', which acquired a few specialized features independently of other choristoderes. In the analyses of Dong and colleagues (2020) it was recovered in a more basal position, outside the clade containing Neochoristodera + the newly named "Allochoristoderes", and in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
with the newly named '' Heishanosaurus'':


References

Choristodera Prehistoric animals of China Fossil taxa described in 2019 Prehistoric reptile genera {{jurassic-reptile-stub