Sunosuchus Phuwiangensis
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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 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, and Thailand and are Jurassic 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 ''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. For example, there are wide pits on the back of the frontal bone. The frontal bone also has a distinctive ridge along part of its midline. The lower jaw has a long symphysis where the two halves come together. This symphysis is formed mostly from the mandibles, but also partially by the splenials. Unlike other goniopholidids, the squamosal bone (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 Junggar Basin in Xinjiang, 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 and tibia, a few metatarsals, and ventral and neck osteoderms. 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 in Sichuan, China. Skulls were collected from the Dashanpu Dinosaur Quarry in 1983, having been found in the Lower Shaximiao Formation. ''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
exoccipital The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cer ...
s 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, 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. This single specimen is the most well preserved vertebrate fossil that has been found from the formation; other vertebrates, including dinosaurs, 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 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 Mongolian 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 ...
turtles, indicating freshwater environments across the region. Marine hybodont sharks and
dipnoan Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, in ...
s 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