Sinosaurus Maxilla
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''Sinosaurus'' (meaning "Chinese lizard") is an extinct genus 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 ...
which lived during the Early
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 Mya. The J ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
. It was a
biped Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
al
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
approximately in length and in body mass. Fossils of the animal were found at the
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 ar ...
, in the
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, 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 ...
Province of
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
.


Discovery and naming

The composite term ''Sinosaurus'' comes from ''Sinae'', the Latin word for the Chinese, and the Greek word ' () meaning "lizard"; thus "Chinese lizard". The specific name, ''triassicus'', refers to the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, the period that the fossils were originally thought to date from. ''Sinosaurus'' was described and named by Chung Chien Young, who is known as the 'Father of Chinese Vertebrate Paleontology', in
1940 A calendar from 1940 according to the Gregorian calendar, factoring in the dates of Easter and related holidays, cannot be used again until the year 5280. Events Below, the events of World War II have the "WWII" prefix. January *January ...
. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, IVPP V34, was found in 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 ar ...
, and consists of two
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 ...
ry (upper jaw) fragments, four maxillary teeth, and a lower jaw fragment with three teeth. The teeth are laterally compressed, and feature fine serrations both at their anterior and posterior edges. The teeth are also variable in size and are curved backwards. This material is too fragmentary to determine the length and weight of this dinosaur. Over the years, other fossils were referred to ''Sinosaurus'', some of which were material that was shown to belong to two
sauropodomorphs 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 ...
. The fossils include a postcrania, with a
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
with three preserved sacral vertebrae. The material assigned to "''Sinosaurus'' postcrania" includes a mix of plateosaurid and melanorosaurid elements. All the material from the Red Beds block has now been reassigned to ''
Jingshanosaurus ''Jingshanosaurus'' (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period 201.3 million years ago that went extinct 199.3 million years ago in the Hettangian Age. Its maximum weight was around 4.3 t ...
'' KMV 8701 was originally discovered in 1987. The specimen was identified as a new species, and was named ''Dilophosaurus sinensis''. Then in 1994, during a field expedition, a more complete specimen was found, and was assigned to the same species. In 2003,
Dong Zhiming Dong Zhiming (Chinese: 董枝明, Pinyin: ''Dǒng Zhimíng''; born January 1937) is a Chinese vertebrate paleontologist formerly employed at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) in Beijing. He began working at the ...
studied the material of ''Sinosaurus triassicus'', finding it to be quite similar to ''Dilophosaurus sinensis''. As ''Sinosaurus'' was named earlier, "Dilophosaurus" ''sinensis'' became its junior synonym. In 2013, a study by Currie et al., confirmed that ''D. sinensis'' was the same animal as ''S. triassicus''Currie, Xing, Wu and Dong, in prep. "Anatomy and relationships of Sinosaurus triassicus ("Dilophosaurus sinensis") from the Lufeng Formation (Lower Jurassic) of Yunnan, China". On the other hand, Wang ''et al.'' (2017) stated that it needs to be further investigated whether ''D. sinensis'' is indeed a junior synonym of ''S. triassicus'', and noted that the two species are different at least in the anatomy of the premaxilla. The authors tentatively assigned ''D. sinensis'' to the genus ''Sinosaurus'', but retained it as a species distinct from ''Sinosaurus triassicus''. Specimen KMV 8701 consists of a skull (measuring 525 mm), and is nearly complete. Dong claimed that animal was about long. It has been assigned now to ''Sinosaurus'', but the specimen still lacks sufficient description and preparation. Over the years, paleontologists referred additional specimens to ''D. sinensis'' which are now assigned to ''Sinosaurus''. Dong (2003) referred specimen LDM-LCA10 which consists of a skull and an incomplete skeleton. In 2012, Xing referred two individuals, ZLJ0003 which consists of a partial skull and an incomplete skeleton, and ZLJT01 which is a juvenile individual that consists of a premaxillary fragment, an incomplete maxilla, a maxillary fragment, a lacrimal, both frontals, both parietals, an incomplete braincase, an incomplete dentary, an atlantal intercentrum, two dorsal rib fragments, and a partial proximal caudal neural arch, to ''Sinosaurus''. In 2012, a new specimen of ''Sinosaurus'' was described, and was found to represent a new species. The species '' Shuangbaisaurus anlongbaoensis'', discovered and named in 2017, has also been considered a synonym of ''Sinosaurus triassicus''.


Description

''Sinosaurus'' was a relatively large theropod during the Early Jurassic, reaching in length and in body mass. According to Carrano ''et al.'' (2012) ''D. sinensis'', now considered to be at least congeneric with ''Sinosaurus triassicus'', can be distinguished based on the fact that a vertical groove is present on the lateral
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 b ...
adjacent to contact with the maxilla. ''Sinosaurus'' is the only "dilophosaurid" known from a complete braincase. ''Cryolophosaurus'', ''Dilophosaurus'', ''
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 Ischigu ...
'' and ''
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 originally as ...
'' are all known from partial braincases. Two partial braincases were found before 2012, and are probably mostly complete, except that large sections are obscured by sediments. In 2011, an exceptionally well-preserved braincase was found, only missing 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, par ...
s and
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 ...
osphenoid.


Classification

Originally thought to be a
coelophysoid Coelophysoidea were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the ...
related to ''
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 preserved ...
'' and ''Cryolophosaurus'', Oliver Rauhut in 2003 showed ''Sinosaurus'' to be a more advanced theropod, related to ''
Cryolophosaurus ''Cryolophosaurus'' ( or ; "CRY-oh-loaf-oh-SAWR-us") is a genus of large theropod dinosaur known from only a single species ''Cryolophosaurus ellioti'', from the early Jurassic of Antarctica. It was one of the largest theropods of the Early Jura ...
'' and "Dilophosaurus" ''sinensis''. In 2013, in an unpublished work, Carano agreed that ''Sinosaurus'' is a theropod. ''Sinosaurus'' has been considered a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
'' in a few works, although now that "Dilophosaurus" ''sinensis'' is referred to it, it is considered valid. ''Dilophosaurus sinensis'' was shown to be a junior synonym of ''Sinosaurus'' in 2003. It is possibly closer to the
Antarctic The Antarctic ( or , American English also or ; commonly ) is a polar region around Earth's South Pole, opposite the Arctic region around the North Pole. The Antarctic comprises the continent of Antarctica, the Kerguelen Plateau and other ...
theropod ''Cryolophosaurus'', based on the fact that the anterior end 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 ...
does not participate in the internal
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, birds ...
and that the maxillary tooth row is completely in front of the eye socket. ''D. sinensis'' was exhibited in 1998 at Dinofest in Philadelphia.Glut, D. F. (1999). Dinosaurs, the Encyclopedia, Supplement 1: McFarland & Company, Inc., 442pp. Although the skull of ''D. sinensis'' sports large nasolacrimal crests superficially like those reconstructed in ''D. wetherilli'', features elsewhere in the skeleton suggest it is closer to
tetanuran Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans ar ...
theropods. Rauhut (2003) regarded ''D. sinensis'' as a basal tetanuran most closely related to ''Sinosaurus'' and ''Cryolophosaurus''. Lamanna ''et al.'' (1998b) examined the material ascribed to ''D. sinensis'' and found it to be synonymous with ''Sinosaurus triassicus''.Lamanna, M. C., Holtz, T. R. Jr, and Dodson, P., 1998, A reassessment of the Chinese Theropod Dinosaur Dilophosaurus sinensis: Journal of Vertebrate Paleontology, Volume 18, Supplement to Number 3. Abstracts of papers. Fifty-eighth annual meeting, Society of Vertebrate Paleontology, Snowbird Ski and Summer Resort, Snowbird, Utah, September 30 – October 3, 1998, p. 57a. This cladistic finding was confirmed in 2003 by Dong. The Lufeng Dinosaurian Museum discovered a new specimen of ''Sinosaurus'' (ZLJT01) in 2007 from the Lufeng Basin. It consists of an incomplete skull and other postcranial fragments. Phylogenetic analysis of this specimen, demonstrates that ''Sinosaurus'' is a more derived theropod, and is not the most basal dilophosaurid, as held by Smith et al. A cladogram was identified by Christophe Hendrickx and
Octávio Mateus Octávio Mateus (born 1975) is a Portuguese dinosaur paleontologist and biologist Professor of Paleontology at the Faculdade de Ciências e Tecnologia da Universidade Nova de Lisboa. He graduated in Universidade de Évora and received his PhD at U ...
. It placed ''Sinosaurus'' and ''Cryolophosaurus'' in a polytomy at the base of Tetanurae. Recent studies placed ''Sinosaurus'' outside the Ceratosauria+Tetanurae clade, while Wang ''et al''. (2016) considered it the basalmost ceratosaur.


Paleobiology


Crest function

''Sinosaurus'' and ''Dilophosaurus'' both possess dual crests. However, it was found that the crests could not be used in combat.


Feeding

The skull of ''Sinosaurus'' has a deep notch between the premaxilla and maxilla. Dong (2003) proposed that the notch was used to house jaw muscles, giving ''Sinosaurus'' a powerful bite. Based on the estimated power of its jaws, ''Sinosaurus'' might have either been a carnivore or a scavenger. Dong suspected that the premaxilla was covered in a narrow, hooked beak, that was used to rip open skin and abdominal flesh. He also thought that the crest would have been used to hold open the abdominal cavity while feeding. Dong studied the feet of ''Sinosaurus'' as well, finding a resemblance with the feet of modern vultures. The feet of ''Sinosaurus'' were probably adapted to help it feed on large-bodied animals, such as prosauropods.


Paleopathology

A study by Xing ''et al.'' (2013) examined the effect of the traumatic loss of teeth on the dental alveolus (the socket in the jaw where the roots of teeth are held) in dinosaurs. ''Sinosaurus'' is the first dinosaur where remodeling of the alveolus in the jaw was observed. The authors concluded that this finding "contributes to mounting evidence suggesting theropods were highly resilient to a broad spectrum of traumas and diseases." The dental alveolus found on ''Sinosaurus'' is the first documented dental pathology found on a dinosaur.


Paleoecology


Provenance and occurrence

The type specimen of ''Sinosaurus triassicus'' IVPP V34 was recovered in the Zhangjiawa Member of the Lufeng Formation, in Yunnan, China. These remains were discovered at the Dark Red Beds that were deposited during the
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and is ...
stage of the Jurassic period, approximately 196-183 million years ago. Several other discoveries referred to ''Sinosaurus'' were made in the Zhangjiawa Member: specimens IVPP V97 (postcrania), IVPP V36 (teeth), IVPP 37 (teeth), IVPP V88 (ilium), IVPP V35 (teeth and postcranial bones), IVPP V100 and IVPP V48 (teeth and postcranial bones) discovered in 1938 by M. Bien & C.C. Young,M. N. Bien. 1940. Discovery of Triassic saurischian and primitive mammalian remains at Lufeng, Yunnan. Bulletin of the Geological Society of China 20(3/4):225-234 FMNH CUP 2001–2003 discovered by E. Oehler and Hu. Specimens FMNH CUP 2097, FMNH CUP 2098, FMNH CUP 2004, FMNH CUP 2005 were discovered in 1948 by M. Bien & C.C. Young at Zhangjiawa Member, as well. ''Sinosaurus'' sp. fossils have been found in the Zhenzhuchong Formation, and were previously thought to be a poposaur, although they might have only been from the equivalent Lufeng Formation. Specimen IVPP V504, referred to ''Sinosaurus'', a maxilla with four teeth, was collected by Lee in the 1940s, in the Dull Purplish Beds of Shawan Member of the Lufeng Formation, that were deposited during the
Hettangian The Hettangian is the earliest age and lowest stage of the 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 (My ...
stage of the Jurassic period, approximately 201-199 million years ago. Several other discoveries were made in the Shawan Member: parts of two skeletons attributed to ''Sinosaurus'' were discovered by Sou in 1956,C.-C. Young. 1966. On a new locality of the Lufengosaurus of Yunnan. Vertebrata PalAsiatica 10(1):64–67 specimen IVPP V279 (tooth) was discovered by C.C. Young in 1938, in dark red clayish sandstone, and specimen IVPP V381 (several teeth) was discovered by C.C. Young, in blue mudstone. The ''D. sinensis'' remains, KMV 8701, a nearly complete skeleton, now referred to ''Sinosaurus'', were recovered in the Shawan Member of Lufeng Formation. This material was discovered in 1987 in the Dull Purplish Beds that were deposited during the Hettangian stage of the Early Jurassic, approximately 201-199 million years ago.


Fauna and habitat

In the
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 ar ...
, ''Sinosaurus'' shared its
paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
with
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
like ''
Morganucodon ''Morganucodon'' (" Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represe ...
'', ''
Oligokyphus ''Oligokyphus'' ("few cusps") is an extinct genus of advanced herbivorous cynodonts of the late Triassic to early Jurassic periods. Originally considered to be an early mammal, it is now classified as a Mammaliamorph (nearly a mammal) because ''O ...
'', and '' Bienotherium'';
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
s like ''
Pachysuchus ''Pachysuchus'' is a Nomen dubium, dubious extinct genus of basal sauropodomorph dinosaur from the Early Jurassic of China. ''Pachysuchus'' is known from a poorly preserved partial Rostrum (anatomy), rostrum that was described from the Lower Luf ...
'';
diapsids Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ag ...
like '' Strigosuchus'';
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 ...
like '' Platyognathus'' and ''
Microchampsa ''Microchampsa'' is an extinct genus of protosuchian crocodylomorph that existed during the Early Jurassic. Fossils have been found from stratum 6 of the Dahuangtian locality, an outcrop of the Lower Red Beds of the Lufeng Formation in Yunnan, C ...
''; the early mammal ''
Hadrocodium ''Hadrocodium wui'' is an extinct mammaliaform that lived during the Sinemurian stage of the Early Jurassic approximately in the Lufeng Formation of the Lufeng Basin in what is now the Yunnan province in south-western China (, paleocoordinates ...
''; and other early reptiles. Contemporary dinosaurs include indeterminate sauropods; the early
thyreophora Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous. Thyreophorans are characterized by the presence of body ...
ns '' Bienosaurus lufengensis'' and '' Tatisaurus oehleri''; the supposed chimeric ornithopod " Dianchungosaurus lufengensis"; the
prosauropods Plateosauridae is a family (biology), family of plateosaurian sauropodomorphs from the Late Triassic of Europe, Greenland, Africa and Asia. Although several dinosaurs have been classified as plateosaurids over the years, the family Plateosaurida ...
'' Gyposaurus sinensis'', '' Lufengosaurus huenei'', '' L. magnus'', '' Jingshanosaurus xinwaiensis'',Y. Zhang, and Z. Yang. (1995). A new complete osteology of Prosauropoda in Lufeng Basin, Yunnan, China. Yunnan Publishing House of Science and Technology, Kunming, China 1–100. hinese/ref> '' Kunmingosaurus wudingensis'', '' Chinshakiangosaurus chunghoensis'', '' Yunnanosaurus huangi'', "Y." ''robustus'', and an unnamed taxon; and the theropods ''
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 ...
'', ''
Eshanosaurus ''Eshanosaurus'' is a genus of a dinosaur from the early Jurassic Period. It is known only from a fossil partial lower jawbone, found in China. It may be a therizinosaurian, and if so the earliest known coelurosaur. Discovery and naming The typ ...
'', and ''
Coelophysis ''Coelophysis'' ( traditionally; or , as heard more commonly in recent decades) is an extinct genus of coelophysid theropod dinosaur that lived approximately 228 to 201.3 million years ago during the latter part of the Triassic Period from t ...
'' sp. '' Changpeipus'' footprints have been found in the Lufeng Formation. In 2009, a study led by Li-Da Xing found that footprints from the Lufeng Formation were unique among ichnogenera, and named the footprints ''Changpeipus pareschequier''. The study hypothesized that they were produced by a
coelophysoid Coelophysoidea were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the ...
; there are many possible trackmakers, however, including both ''Sinosaurus'' and ''Coelophysis'' sp.


References

{{Taxonbar, from=Q149083 Early Jurassic dinosaurs of Asia Prehistoric neotheropods Fossil taxa described in 1940 Taxa named by Yang Zhongjian Paleontology in Yunnan