Litorosuchus
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''Litorosuchus'' is a genus of armored, semiaquatic archosauriform reptile 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 ...
of China, closely related to the morphologically similar '' Vancleavea''. It contains one species, ''L. somnii''.


Description

For an archosauriform, ''Litorosuchus'' was medium-sized at some in length. It shows clear adaptations to a semiaquatic lifestyle: the nasal opening is retracted and angled upwards, the tail is tall and long (about 60% of the animal's entire length), the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...
is short and broad, the feet are webbed (as shown by fossilized skin impressions), the neck is long and slender, and the snout is long, with many of the teeth being conical. In many of these ways, ''Litorosuchus'' resembles the similarly semiaquatic '' Vancleavea'' from North America. Like ''Vancleavea'', the body of ''Litorosuchus'' was covered in bony plates known as
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 (extinc ...
s, of which eight distinct types can be observed: two rows of rectangular-to-roundish semi-concave osteoderms with central ridges covered the neck and back; more oval-shaped osteoderms, also with ridges, covered the sides of the trunk and the rest of the neck; a row of tall, spike-like osteoderms lined the top of the tail (similar to ''Vancleavea''); small, thin, and variably-shaped osteoderms covered the arm above the hand and the foot above the ankle; sheets of tiny osteoderms covered the hands and feet; rows of broad, oval-shaped osteoderms covered the bottom of the trunk; similarly broad but semi-concave osteoderms covered the bottom portion of the tail; and oval, keeled osteoderms with a notch on the back edge covered the remainder of the tail. Out of these eight types, the former four are found in other archosauriforms, but the latter four are entirely unique to ''Litorosuchus''. Asides from the aforementioned characteristics, ''Litorosuchus'' can also be differentiated from other archosauriforms by a unique combination of other characters: only two teeth are present on 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 are situated close to the front of the bone; the
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 ...
ry
process A process is a series or set of activities that interact to produce a result; it may occur once-only or be recurrent or periodic. Things called a process include: Business and management *Business process, activities that produce a specific se ...
, on the back edge of the premaxilla, is relatively long; the nasal process of the maxilla extends behind the back edge of the nostril; there is a large, canine-like (caniniform) tooth on each maxilla (also seen in ''Vancleavea''); the
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 modern a ...
is T-shaped and extends down to the
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 ...
, separating the latter from the eye socket; the width of the skull roof between 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 ...
e is very thin, being only one-fifth the width of the skull roof between the eye sockets; a vertical ridge is present on the sides of most of the caudal vertebrae; and the joint between the
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
and the
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. ...
is a simple
butt joint A butt joint is a technique in which two pieces of material are joined by simply placing their ends together without any special shaping. The name "butt joint" comes from the way the material is joined. The butt joint is the simplest joint to ma ...
(the bones meet in flat surfaces, not ball-and-socket joints).


Discovery and naming

''Litorosuchus'' is known from one nearly-complete skeleton embedded in a slab of
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
and lying on its left side. The rocks in which the specimen was found are located in
Fuyuan County, Yunnan Fuyuan () is under the administration of Qujing City, in the east of Yunnan province, China, bordering Guizhou province to the east. Administrative divisions Fuyuan County has 2 subdistricts, 9 towns and 1 ethnic township. ;2 subdistricts * Zho ...
, and belong to the Zhuganpo Member of the
Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic ...
-age Falang Formation. The specimen is stored in 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 sugges ...
under the number IVPP V 16978. The genus name is derived from Latin ''litoralis'' ("littoral"), in reference to the habitat of this animal, and Greek ''soukhos'' ("crocodile"). The species name, ''somnii'', comes from the Latin ''somnium'' ("dream"); this is because the lead author dreamt of the animal while trying to come up with a name for it.


Classification

In 2016, ''Litorosuchus'' was found to be closely related to '' Vancleavea'' in a phylogenetic analysis, with both being non- archosaur archosauriforms. The results of said analysis are reproduced partially below.


Paleoecology

The Zhuganpo Member of the Falang Formation, from which ''Litorosuchus'' is known, would have been a coastal, shallow-water environment. In addition to various actinopterygian fish, various reptiles are known from this location; they include the terrestrial '' Macrocnemus'' as well as the aquatic ''
Lariosaurus ''Lariosaurus'' is an extinct genus of nothosaurid from the Middle Triassic (late Anisian to late Ladinian stage) of central and western Europe and China. With a length of just , it was one of the smallest known nothosaurs. First discovered at Pe ...
'', ''
Keichousaurus ''Keichousaurus'' (key-cho-saurus) is a genus of marine reptile in the pachypleurosaur family which went extinct at the close of the Triassic in the Triassic-Jurassic extinction event. The name derives from Kweichow (now Guizhou Province) in ...
'', '' Yunguisaurus'', ''
Qianxisaurus ''Qianxisaurus'' is an extinct genus of pachypleurosaur or alternatively a basal eosauropterygian known from the Middle Triassic (Ladinian age) of Guizhou Province, southwestern China. It contains a single species, ''Qianxisaurus chajiangens ...
'', ''
Tanystropheus ''Tanystropheus'' (Greek ~ 'long' + 'hinged') is an extinct archosauromorph reptile from the Middle and Late Triassic epochs. It is recognisable by its extremely elongated neck, which measured long—longer than its body and tail combined. T ...
'', '' Anshunsaurus'', and ''
Glyphoderma ''Glyphoderma'' is an extinct genus of placodont reptile from the Middle Triassic of China. It differs from its relative '' Psephochelys'' in having three, rather than one, fused osteoderms on the posterior skull surface, and has an earlier tempo ...
''. Besides ''Litorosuchus'' and '' Vancleavea'', other possibly semiaquatic archosauriforms include ''
Qianosuchus ''Qianosuchus'' is an extinct genus of aquatic poposauroid archosaur from the middle Triassic (Anisian) Guanling Formation of Pan County, China. It is represented by two nearly complete skeletons and a crushed skull preserved in the limestone. ...
'' and ''
Diandongosuchus ''Diandongosuchus'' is an extinct genus of archosauriform reptile, possibly a member of the Phytosauria, known from the Middle Triassic of China. The type species ''Diandongosuchus fuyuanensis'' was named in 2012 from the Falang Formation of Y ...
''; both show some of the morphological specializations associated with semiaquatic lifestyles, and the latter was found with fish bones in its stomach. In addition,
phytosaur Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent g ...
s and proterochampsids also have retracted nostrils. The large number of independent acquisitions of semiaquatic lifestyles among archosauriforms may be indicative of plasticity in the lifestyles of these animals, which could also explain the prevalence of semiaquatic crocodiles and birds in the modern world.


References

{{Taxonbar, from=Q28432480 Prehistoric reptile genera Prehistoric archosauriforms Middle Triassic reptiles of Asia Triassic China Fossils of China Fossil taxa described in 2016 Paleontology in Yunnan