Sinosaurosphargis Yunguiensis
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''Sinosaurosphargis'' 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 basal
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
saurosphargid Saurosphargidae is an extinct family of marine reptiles known from the early Middle Triassic (Anisian stage) of Europe and China. The type species of the family is '' Saurosphargis volzi'', named by Friedrich von Huene in 1936 based on a single ...
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
known from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
(
Anisian In the geologic timescale, the Anisian is the lower stage or earliest age of the Middle Triassic series or epoch and lasted from million years ago until million years ago. The Anisian Age succeeds the Olenekian Age (part of the Lower Triassic Ep ...
age)
Guanling Formation The Guanling Formation is a Middle Triassic (Anisian or Pelsonian in the regional chronostratigraphy) geologic formation in southwestern China. Description The formation encompasses two members. The first member is primarily calcareous mudston ...
of
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 ...
and
Guizhou Guizhou (; formerly Kweichow) is a landlocked province in the southwest region of the People's Republic of China. Its capital and largest city is Guiyang, in the center of the province. Guizhou borders the autonomous region of Guangxi to t ...
Provinces, southwestern
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 ...
. It contains a single species, ''Sinosaurosphargis yunguiensis''.


Discovery

''Saurosphargis'' is known from several individuals, all of which were collected from Member II of the
Guanling Formation The Guanling Formation is a Middle Triassic (Anisian or Pelsonian in the regional chronostratigraphy) geologic formation in southwestern China. Description The formation encompasses two members. The first member is primarily calcareous mudston ...
, dating to the Pelsonian substage of the latest
Anisian In the geologic timescale, the Anisian is the lower stage or earliest age of the Middle Triassic series or epoch and lasted from million years ago until million years ago. The Anisian Age succeeds the Olenekian Age (part of the Lower Triassic Ep ...
stage of the early
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
, about 243
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. 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 V 17040 and
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
IVPP V 16076 are housed at the
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
in Beijing, and represent a nearly complete articulated
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
and
skull 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 a partial disarticulated
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l skeleton including dorsal
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
,
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s, with
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 and gastral rib fragments, respectively. ZMNH M 8797, an incomplete
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l skeleton showing a very well preserved right
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
housed at the Zhejiang Museum of Nature History of Hangzhou, was also referred to ''Saurosphargis'' in its original description. The holotype and ZMNH M 8797 were collected at Yangmazhai of
Luoping County Luoping County () is located in Qujing City, in eastern Yunnan province, China, bordering Guizhou province to the east and the Guangxi Zhuang Autonomous Region to the southeast. The county of Luoping in eastern Yunnan is noted for its springtim ...
, Yunnan Province, while the paratype came from Yangjian of
Pan County Panzhou () is a county-level city in southwestern Guizhou province, China, on the border with Yunnan province to the west. It is under the administration of the prefecture-level city of Liupanshui. Administrative divisions As of 2017, Panzhou is ...
, Guizhou Province. An additional uncatalogued LPV specimen was also referred to ''S. yunguiensis''. These specimens all came from the
Lagerstätte A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
discovered during a 2007 geological mapping project with a diverse record of marine life called Luoping Biota, that yielded, apart from ''Sinosaurosphargis'', various invertebrates, fish, basal
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
s, ''
Atopodentatus ''Atopodentatus'' is an extinct genus of basal sauropterygian known from the early Middle Triassic (Pelsonian substage, Anisian to Ladinian stage) of Luoping County, Yunnan Province, southwestern China. It contains a single species, ''Atopodenta ...
'', the basal saurosphargid ''
Largocephalosaurus ''Largocephalosaurus'' is an extinct genus of basal saurosphargid, a marine reptile known from the Middle Triassic (Anisian age) Guanling Formation of Yunnan and Guizhou Provinces, southwestern China. It contains a type species, ''Largocephalo ...
'', and several species of
eosauropterygia Sauropterygia (" lizard flippers") is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria bec ...
n, including both
pachypleurosaur left, 220px, '' Pachypleurosaurus'' Pachypleurosauria is an extinct clade of primitive sauropterygian reptiles that vaguely resembled aquatic lizards, and were limited to the Triassic period. They were elongate animals, ranging in size from , w ...
s and
nothosaurid Nothosauridae are an extinct family of carnivorous aquatic sauropterygian reptiles from the Triassic time period of China, France, Germany, Israel, Italy, Netherlands, Russia, Switzerland, and northern Africa. Phylogeny The cladogram shown bel ...
s.


Etymology

''Sinosaurosphargis'' was first described and named by Chun Li, Olivier Rieppel, Xiao-Chun Wu, Li-Jun Zhao and Li-Ting Wang in
2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrate ...
and 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 ...
is ''Sinosaurosphargis yunguiensis''. The generic name is derived from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''sino'', meaning "China", ''sauros'', meaning "lizard", and ''sphargis'', the old genus name for the
leatherback turtle The leatherback sea turtle (''Dermochelys coriacea''), sometimes called the lute turtle or leathery turtle or simply the luth, is the largest of all living turtles and the heaviest non-crocodilian reptile, reaching lengths of up to and weights ...
, (in reference to the many similarities ''Sinosaurosphargis'' shares with ''
Saurosphargis ''Saurosphargis'' is an extinct genus of a basal marine reptile, saurosphargid, known from the Middle Triassic (Anisian age) of southwestern Poland and eastern Netherlands. It contains a single species, ''Saurosphargis volzi''. Discovery ' ...
'', a taxon whose holotype was lost and was thus considered a mystery), following the pattern of ''
Sinosauropteryx ''Sinosauropteryx'' (meaning "Chinese reptilian wing", ) is a compsognathid dinosaur. Described in 1996, it was the first dinosaur taxon outside of Avialae (birds and their immediate relatives) to be found with evidence of feathers. It was covere ...
''. Like the name of ''Saurosphargis'' itself, it also refers to ''Sinosaurosphargis dorsal
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 ...
"body armor" and broadened ribs forming a closed chest rib basket - traits which are seemingly transitional between
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 and other reptiles. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
''yunguiensis'' is derived from the names of the neighboring
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 ...
and
Guizhou Guizhou (; formerly Kweichow) is a landlocked province in the southwest region of the People's Republic of China. Its capital and largest city is Guiyang, in the center of the province. Guizhou borders the autonomous region of Guangxi to t ...
Provinces, where the fossils were found.


Description

''Sinosaurosphargis'' was a small reptile, measuring long without the missing tail. It is a basal
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. The earliest marine reptile mesosaurus (not to be confused with mosasaurus), arose in the Permian period during the ...
, closely related to ''Saurosphargis'' from the Lower Muschelkalk of southwestern
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...
and eastern
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
. ''Sinosaurosphargis'' possesses retracted external
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
s, and shows closed upper
temporal 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 a ventrally open cheek. Its trunk region is short and rounded, and covered by a shield composed of small
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, just like its neck and the proximal portions of all four limbs. Its back vertebrae show elongated, distally expanded transverse projections. The chest
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s are broadened and flat, contacting one another along their length, and forming a closed chest rib basket. Li ''et al.'' (2014) found some of these traits to be shared with all
saurosphargid Saurosphargidae is an extinct family of marine reptiles known from the early Middle Triassic (Anisian stage) of Europe and China. The type species of the family is '' Saurosphargis volzi'', named by Friedrich von Huene in 1936 based on a single ...
s, while others are exclusive to ''Saurosphargis'' and ''Sinosaurosphargis''. These genera have broader and flatter rib baskets than basal saurosphargids like ''
Largocephalosaurus ''Largocephalosaurus'' is an extinct genus of basal saurosphargid, a marine reptile known from the Middle Triassic (Anisian age) Guanling Formation of Yunnan and Guizhou Provinces, southwestern China. It contains a type species, ''Largocephalo ...
'', and also transverse projections of back vertebrae and proximal portions of chest ribs that are much more robust, being much wider than the spacing between them.


Phylogeny

The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
is simplified after the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of Li ''et al.'' (2011) and shows the placement of ''Sinosaurosphargis'' within
Sauria Sauria is the clade containing the most recent common ancestor of archosaurs (such as crocodilians, dinosaurs, etc.) and lepidosaurs ( lizards and kin), and all its descendants. Since most molecular phylogenies recover turtles as more closely re ...
.
Ichthyopterygia Ichthyopterygia ("fish flippers") was a designation introduced by Sir Richard Owen in 1840 to designate the Jurassic ichthyosaurs that were known at the time, but the term is now used more often for both true Ichthyosauria and their more primitiv ...
was removed from the tree, as it caused a less resolved topology. ''Saurosphargis'' was coded solely based on the holotype, as
the Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
material remains unprepared. Li ''et al.'' (2014) presented an updated version of the analysis above, showing interrelationships of all known
Saurosphargidae Saurosphargidae is an extinct family of marine reptiles known from the early Middle Triassic (Anisian stage) of Europe and China. The type species of the family is '' Saurosphargis volzi'', named by Friedrich von Huene in 1936 based on a single ...
species. The removal / inclusion of
Ichthyopterygia Ichthyopterygia ("fish flippers") was a designation introduced by Sir Richard Owen in 1840 to designate the Jurassic ichthyosaurs that were known at the time, but the term is now used more often for both true Ichthyosauria and their more primitiv ...
was found to affect the
topology In mathematics, topology (from the Greek language, Greek words , and ) is concerned with the properties of a mathematical object, geometric object that are preserved under Continuous function, continuous Deformation theory, deformations, such ...
the most - switching the positions of the ''Eusaurosphargis''+''Helveticosaurus'' and Thalattosauriformes clades, and altering the positions of several taxa within
Eosauropterygia Sauropterygia (" lizard flippers") is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria bec ...
, which are not shown.


References

{{Taxonbar, from=Q3961919 Fossil taxa described in 2011 Middle Triassic reptiles of Asia Fossils of China Anisian life Guanling Formation Saurians Prehistoric reptile genera