Shuangbaisaurus
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''Shuangbaisaurus'' (meaning "Shuangbai reptile") is
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
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
, possibly a junior synonym of '' Sinosaurus.'' It lived in the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
of
Yunnan Province Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, and is represented by a single species, ''S. anlongbaoensis'', known from a partial skull. Like the theropods ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
'' and ''Sinosaurus,'' ''Shuangbaisaurus'' bore a pair of thin, midline crests on its skull. Unusually, these crests extended backwards over the level of the eyes, which, along with the unusual orientation of the
jugal bone 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. Ana ...
, led the describers to name it as a new genus. However, ''Shuangbaisaurus'' also possesses a groove between its
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 ...
and
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. T ...
, a characteristic which has been used to characterize ''Sinosaurus'' as a genus. Among the two morphotypes present within the genus ''Sinosaurus'', ''Shuangbaisaurus'' more closely resembles the morphotype that is variably treated as a distinct species, ''S. sinensis'', in its relatively tall skull.


Description

''Shuangbaisaurus'' had a relatively robust (compared to '' Sinosaurus'') skull, with a length of and a length/height ratio of 2.84, not including the thin paired crests that were present at least above the eye sockets. These crests are incompletely preserved, but appeared to have been formed mostly from the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, ...
, as well as possibly 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 ...
and
prefrontal bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most mo ...
s. The crests may have extended further forward, over the
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 * ...
and
lacrimal bone The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
s. Similar crests are seen in ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
'' and ''Sinosaurus''; however, the crests of ''Sinosaurus'' do not extend over the eyes, thus differentiating it from ''Shuangbaisaurus''. At the tip of the jaw, 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 ...
and
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. T ...
are separated by a groove, which is also a defining characteristic of ''Sinosaurus''. The premaxilla would have been taller than it is long, which is similar to specimen LFGT LDM-L10 of ''Sinosaurus triassicus'' but unlike either ''Dilophosaurus'' or specimen KMV 8701 of ''Sinosaurus sinensis''. Also like LFGT LDM-L10, the bottom edge of the premaxilla is much higher than the maxilla. The two bones are at the same level in KMV 8701, and are angled away from each other in ''Dilophosaurus''. Similar to both LFGT LDM-L10 and KMV 8701, the maxilla bears a near-vertical front margin; it is lower and more pointed in ''Dilophosaurus''. The front portion of the bottom margin of the maxilla is angled such that the first maxillary tooth projects forwards. There is a groove running parallel to the tooth row along the bottom rim of the maxilla. From behind, the maxilla is connected to 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 ...
. The lacrimal is located above their joint, enclosing the fossa that surrounds the
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, bird ...
. There is a ridge directed downwards and backwards on the side of the jugal, like in ''
Zupaysaurus ''Zupaysaurus'' (; "ZOO-pay-SAWR-us") is an extinct genus of early theropod dinosaur living during the Norian stage of the Late Triassic in what is now Argentina. Fossils of the dinosaur were found in the Los Colorados Formation of the Ischigual ...
''; this same ridge is horizontal in ''Sinosaurus'' and ''Dilophosaurus''. The same is also true of the joint between the jugal and
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 ...
. The quadratojugal tapers to a point underneath the infratemporal fenestra, where it is joined at the back to the quadrate. Further above, the lacrimal is joined by the long and slender descending branch of the three-pronged postorbital, which is convex in its upper third. From above, the forward-directed branch of the postorbital is offset from the backward-directed branch, being separated by a small recess. The frontals appeared to have been long and large, forming most of the top of the skull. The antorbital fenestra of ''Shuangbaisaurus'' is large, occupying roughly one third the length of the skull. The back margin of the fenestra slopes backwards, like LFGT LDM-L10 and ''Dilophosaurus'' but unlike KMV 8701. Further behind, 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 ...
is keyhole-shaped, with a longer back rim compared to the front rim. The infratemporal fenestra is roughly trapezoidal, with a long axis oriented upwards and backwards towards the top of the rear of the skull. On the top surface of the skull, the rounded supratemporal fenestra is unusually small, with a diameter equivalent to only about half of the length on the top of the skull behind the level of the orbit.


Discovery and naming

''Shuangbaisaurus'' is known from a single specimen, a partial skull missing most of the top of the snout, stored at the Chuxiong Prefectural Museum in
Chuxiong Yi Autonomous Prefecture Chuxiong Yi Autonomous Prefecture (; Chuxiong Yi script: ,IPA: ; Yi script: ꊉꇑꆑꌠꑼꂰ; Yi Pinyin: wop lup nut su yuop mi) is an autonomous prefecture located in central Yunnan Province, China. Chuxiong has an area of . The capital of t ...
,
Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
under the specimen number CPM C2140ZA245. Additionally, the front half and back portion of the jaw are also associated with the skull. The specimen's snout has also become bent to the left due to deformation during fossilization. It was discovered in layers of purple muddy
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, ...
located in Liuna Village, Anlongbao Town, Shuangbai County, Chuxiong Yi, about south of the well-known Lufeng deposits. These layers have been recognized as belonging to the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
Fengjiahe Formation The Fengjiahe Formation is a geological formation in China. It dates back to the Early Jurassic, most likely to the Pliensbachian. The formation is up to 1500 metres thick and consists of "purple-red mudstone and argillaceous siltstone interbedde ...
, which has been tentatively dated to the
Hettangian The Hettangian is the earliest age and lowest stage of the Jurassic Period of the geologic timescale. It spans the time between 201.3 ± 0.2 Ma and 199.3 ± 0.3 Ma (million years ago). The Hettangian follows the Rhaetian (part of the Triassi ...
. In 2017, the type and only species of the genus ''Shuangbaisaurus'', ''S. anlongbaoensis'', was formally described and named by Wang Guo-Fu, You Hai-Lu, Pan Shi-Gang, and Wang Tao. The generic name refers to Shuangbai County, and the specific name refers to Anlongbao Town (with Anlongbao literally meaning "dragon-placing fort"), and thus both elements of the binomial name reference its provenance. In a 2019 conference abstract, Philip Currie and colleagues suggested that CPM C2140ZA245 fell within the range of variation of specimens assigned to ''Sinosaurus triassicus''.


Classification

After ''Dilophosaurus'' and ''Sinosaurus'', ''Shuangbaisaurus'' is the third large
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
known definitely to have had paired crests on its skull (though the smaller ''
Coelophysis kayentakatae ''Coelophysis''? ''kayentakatae'' is an extinct species of neotheropod dinosaur that lived approximately 200 - 196 million years ago during the early part of the Jurassic Period in what is now the southwestern United States. While originall ...
'' also has similar crests). ''Shuangbaisaurus'' seems to generally be similar to the roughly contemporaneous ''Sinosaurus''. However, they are notably differentiated in that the crests clearly extend over the orbit in ''Shuangbaisaurus'', whereas they are restricted to extensions of the nasal and lacrimal that do not reach past the orbit in ''Sinosaurus''. Similarly, the backward and downward orientation of the jugal is a trait of ''Shuangbaisaurus'' not seen in ''Sinosaurus'', where it is horizontal. However, the vertical groove separating the premaxilla and maxilla, which seems to be present in ''Shuangbaisaurus'', was denoted as a distinguishing characteristic of ''Sinosaurus'' by Matthew Carrano and colleagues in 2012, thus complicating its referral to a new genus by Wang ''et al.'' A further issue in the relation between ''Shuangbaisaurus'' and ''Sinosaurus'' involves the taxonomic convolution that has befallen ''Sinosaurus''. At least one species is recognized in the genus, ''Sinosaurus triassicus'', consisting of specimens LFGT LDM-L10 and LFGT ZLJT01; another specimen, KMV 8701, originally named "Dilophosaurus" ''sinensis'', has been referred to either ''S. triassicus'' or a second species ''S. sinensis''. KMV 8701 approaches ''Shuangbaisaurus'' more closely in its tall skull and premaxilla than specimens referred to ''S. triassicus''. Wang ''et al.'' elected to retain ''S. sinensis'' as a separate species owing to these differences. No phylogenetic analysis has been conducted to quantitatively determine the relationships of ''Shuangbaisaurus''.


Paleoecology

Aside from ''Shuangbaisaurus'', several
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
dinosaurs originate from the Fengjiahe Formation. These include '' Chinshakiangosaurus chunghoensis'' (a possible
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', ' lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their ...
), '' Lufengosaurus sp.'', '' Yimenosaurus youngi'', and ''
Yunnanosaurus huangi ''Yunnanosaurus'' ( ) is an extinct genus of sauropodomorph dinosaur that lived approximately 199 to 183 million years ago in what is now the Yunnan Province, in China, for which it was named. ''Yunnanosaurus'' was a large sized, moderately-buil ...
'' (a juvenile specimen, which was also discovered in Shuangbai County). Ostracods of the genus ''
Darwinula ''Darwinula'' is a genus of ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several or ...
'' have also been found. A diverse
ichnofauna Ichnofauna is the diversity of fauna based on the ichnological ( tracks and footprints) evidences. This term is often used by vertebrate paleontologists.Mateus, O., & Milàn, J. 2010. A diverse Upper Jurassic dinosaur ichnofauna from central‐west ...
is also known from deposits of the Fengjiahe Formation, including the
ichnotaxa An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxa'' comes from the Greek ίχνος, ''ichnos'' meaning ''track'' and ταξις, ''taxis'' meaning ...
'' Eubrontes platypus'', '' Grallator limnosus'', '' Paracoelurosaurichnus monax'', '' Schizograllator xiaohebaensis'', '' Youngichnus xiyangensis'', and '' Zhengichnus jinningensis'' (with the last four sometimes considered synonymous with ''Grallator''). Temporally, the Fengjiahe Formation is equivalent to the
Lower Lufeng Formation The Lufeng Formation (formerly Lower Lufeng Series) is a Lower Jurassic sedimentary rock formation found in Yunnan, China. It has two units: the lower Dull Purplish Beds/Shawan Member are of Hettangian age, and Dark Red Beds/Zhangjia'ao Member a ...
, from where ''Sinosaurus'' is known alongside a diverse assemblage consisting of another theropod, ''
Lukousaurus ''Lukousaurus'' is an archosauromorph based on most of a small skull's snout, displaying distinctive lachrymal horns, found in the Early Jurassic-age Lower Lufeng Formation, Yunnan, China and was described by Chung Chien Young in 1940. The ge ...
''; sauropodmorphs (including ''Chinshakiangosaurus sp.'', ''Lufengosaurus huenei'', ''L. magnus'', '' Xingxiulong chengi'' and ''Yunnanosaurus huangi'');
ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
ns;
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
;
cynodont The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide varie ...
s and mammals; turtles; amphibians; and bivalves.


See also

*
2017 in archosaur paleontology The year 2017 in archosaur paleontology was eventful. Archosaurs include the only living dinosaur group — birds — and the reptile crocodilians, plus all extinct dinosaurs, extinct crocodilian relatives, and pterosaurs. Archosaur palaeontolog ...


References

{{Taxonbar, from=Q29051703 Prehistoric theropods Early Jurassic dinosaurs of Asia Hettangian life Jurassic China Fossils of China Paleontology in Yunnan Fossil taxa described in 2017