Sunosuchus Miaoi
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''Sunosuchus'' 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
goniopholidid Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle Ju ...
mesoeucrocodylia Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day. Diagnosis It was long known that M ...
n. Fossils are known from
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 ...
,
Kyrgyzstan Kyrgyzstan,, pronounced or the Kyrgyz Republic, is a landlocked country in Central Asia. Kyrgyzstan is bordered by Kazakhstan to the north, Uzbekistan to the west, Tajikistan to the south, and the People's Republic of China to the east. ...
, and
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
and are
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 ...
in age, although some may be
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 ...
. Four species are currently assigned to the genus: 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 ...
''S. miaoi'' and the species ''S. junggarensis'', ''S. shartegensis'', and ''S. shunanensis''. All species are from China. ''
Goniopholis ''Goniopholis'' (meaning "angled scale") is an extinct genus of goniopholidid crocodyliform that lived in Europe and Africa during the Late Jurassic and Early Cretaceous. Being semi-aquatic it is very similar to modern crocodiles. It ranged from ...
phuwiangensis'', also from Thailand, was reassigned to ''Sunosuchus'' by Andrade ''et al.'' (2011). The material from Kyrgyzstan has not been assigned to any species.


Description

''Sunosuchus'' has a long, narrow snout and a small
skull table The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
. Several characters help diagnose ''Sunosuchus'' and distinguish it from other
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
. For example, there are wide pits on the back of 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 ...
. 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 ...
also has a distinctive ridge along part of its midline. The lower jaw has a long
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...
where the two halves come together. This symphysis is formed mostly from the
mandible 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 tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s, but also partially by the
splenial The splenial is a small bone in the lower jaw of reptiles, amphibians and birds, usually located on the lingual side (closest to the tongue) between the angular and surangular The suprangular or surangular is a jaw bone found in most land verteb ...
s. Unlike other goniopholidids, 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 co ...
(which is found near the back of the skull) is narrow.


Species

250px, Holotype of ''S. shartegensis'', which may have instead belonged to the related genus '' Chalawan'', alt=, left ''S. junggarensis'' is the best known species of ''Sunosuchus''. It was first described in 1996 from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
Junggar Basin The Junggar Basin () is one of the largest sedimentary basins in Northwest China. It is located in Xinjiang, and enclosed by the Tarbagatai Mountains of Kazakhstan in the northwest, the Altai Mountains of Mongolia in the northeast, and the Heaven ...
in
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
, China. Material belonging to ''Sunosuchus'' was collected from Kyrgyzstan in the 1980s and was described in 2000. Many teeth were found, as well as a dorsal vertebra and some vertebral centra, some pelvic bones, part of a
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...
and
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, a few metatarsals, and ventral and neck
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s. The only parts of the skull uncovered were squamosal bones. The squamosals are similar to those of other species of ''Sunosuchus'' but not those of other genera, which indicates that the material belongs to a species of ''Sunosuchus''. While the specimens bear a strong resemblance to the bones of ''S. junggarensis'', they have not been assigned to any species. A fifth species, ''S. shunanensis'', was described in 2005 from the Middle Jurassic of
Zigong Zigong (, ), formed by the merger of the two former towns of Ziliujing (Tzuliuching, literally "self-flow well") and Gongjing (Kungching, literally "offering well"), is a prefecture-level city in Sichuan, southwestern China. Demographics Accord ...
in
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
, China. Skulls were collected from the Dashanpu Dinosaur Quarry in 1983, having been found in the
Lower Shaximiao Formation The Shaximiao Formation () is a Middle to Late Jurassic aged geological formation in Sichuan, China, most notable for the wealth of dinosaurs fossils that have been excavated from its strata. The Shaximiao Formation is exposed in and around the ...
. ''S. shunanensis'' has a longer snout than other species, being around three times the length of the postorbital region, or the portion of the skull behind the eyes. It also has a wider skull table than other species. The skull is widest at the back rather than between the eyes, a feature only seen in ''S. shunanensis''. Near the back of each maxilla there is a distinct depression. There are unique ridges across the surface of the skull, one pair at the front of the eye socket on the lacrimals, and a second along the sides of the bacioccipitals and the undersurfaces of the exoccipitals at the base of the skull.


Formerly assigned to ''Sunosuchus''

''Sunosuchus thailandicus'' was described from northeastern Thailand in 1980. It is known only from the
mandible 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 tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
, which is very robust. The tip of the jaw is spoon shaped and wider than the portion of the jaw immediately behind it. The mandible was collected from the
Phu Kradung Formation The Early Cretaceous Phu Kradung Formation is the lowest member of the Mesozoic Khorat Group which outcrops on the Khorat Plateau in Isan, Thailand. This geological formation consists of micaceous, brown to reddish-brown siltstone beds with minor ...
near the town of
Nong Bua Lamphu Nong Bua Lam Phu () is a town in Thailand, capital of Nong Bua Lamphu Province. It is on the central eastern border of the province, approximately 45 kilometers south-west of the city of Udon Thani and from there, accessed by route 210. The town l ...
. This single specimen is the most well preserved vertebrate fossil that has been found from the formation; other vertebrates, including
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 ...
s, are known only from fragmentary remains. The age of the Phu Kradung Formation is uncertain. It was once thought to be Early Jurassic, which would have made ''S. thailandicus'' the oldest species of ''Sunosuchus''. However, the formation has more recently been considered to be Late Jurassic to Early Cretaceous in age, potentially making ''S. thailandicus'' the youngest species of the genus. However, a newly discovered specimen showed that ''S. thailandicus'' is sufficiently distinct from other nominal species of ''Sunosuchus'' to be placed in its own genus, ''
Chalawan ''Krai Thong'' or ''Kraithong'' ( th, ไกรทอง, ) is a Thai folktale, originating from Phichit Province. It tells the story of Chalawan, a crocodile lord who abducts a daughter of a wealthy Phichit man, and Kraithong, a merchant from No ...
''.Martin, J. E.; Lauprasert, K.; Buffetaut, E.; Liard, R.; Suteethorn, V. (2013). "A large pholidosaurid in the Phu Kradung Formation of north-eastern Thailand". In Angielczyk, Kenneth. Palaeontology: n/a. doi:10.1111/pala.12086. edit


Phylogeny

''Sunosuchus'' was initially classified as a pholidosaurid because its jaws were long, making it a longirostrine, or long-snouted, crocodyliform. Most pholidosaurs are longirostrine, while goniopholidids usually have shorter snouts. Despite the similarities with pholidosaurs, ''Sunosuchus'' shares several features with goniopholidids that ally it with the group. Among these features are small
supratemporal fenestrae 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 ...
and openings at the front of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
. The results of a phylogenetic analysis conducted in 2011 focusing on the interrelationships of goniopholidids is shown below.


Paleobiology

The material from Kyrgyzstan is the westernmost record of the geographic range of ''Sunosuchus''. The strata from which this material has been found are similar to those of
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
n and Chinese Middle and Late Jurassic strata. These areas shared a similar fauna that is characterized by crocodyliforms such as ''Sunosuchus'' as well as
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s and
xinjiangchelyid Xinjiangchelyidae is an extinct family of turtles known from the Lower Jurassic to the Middle Cretaceous of Asia and western Europe. They have generally been interpreted as either being basal cryptodires or placed outside of crown Testudines. Ge ...
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s, indicating freshwater environments across the region. Marine
hybodont Hybodontiformes, commonly called hybodonts, are an extinct group of shark-like chondrichthyans, which existed from the late Devonian to the Late Cretaceous. They form the group of Elasmobranchii closest to neoselachians, the clade of modern shark ...
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s and dipnoans are found in the Kyrgyzstan strata, suggesting that the area was
estuarine An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environment ...
at the time rather than entirely freshwater.


References

{{Taxonbar, from=Q3977038 Middle Jurassic crocodylomorphs Jurassic reptiles of Asia Late Jurassic crocodylomorphs Neosuchians Prehistoric pseudosuchian genera