Litorosuchus
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''Litorosuchus'' is a genus of armored, semiaquatic archosauriform
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
of
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, 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 (: 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 either side ...
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 (extinct amph ...
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 mammals h ...
, and are situated close to the front of the bone; the
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
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 s ...
, on the back edge of the premaxilla, is relatively long; the
nasal Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination: * With reference to the human nose: ** Nasal administration, a method of pharmaceutical drug delivery * ...
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 is T-shaped and extends down to the lacrimal bone, 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 Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
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 may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
and the
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel; : calcanei or calcanea) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is t ...
is a simple
butt joint A butt joint is a joinery, wood joint in which the end of a piece of material is simply placed (or “butted”) against another piece. The butt joint is the simplest joint. An unreinforced butt joint is also the weakest joint, as it provides a ...
(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 is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
and lying on its left side. The rocks in which the specimen was found are located in Fuyuan County, Yunnan, and belong to the Zhuganpo Member of the Ladinian-age Falang Formation. The specimen is stored in the Institute of Vertebrate Paleontology and Paleoanthropology 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 Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
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'', '' Keichousaurus'', '' Yunguisaurus'', '' Qianxisaurus'', '' Tanystropheus'', '' Anshunsaurus'', and '' Glyphoderma''. Besides ''Litorosuchus'' and '' Vancleavea'', other possibly semiaquatic archosauriforms include '' Qianosuchus'' and '' Diandongosuchus''; both show some of the morphological specializations associated with semiaquatic lifestyles, and the latter was found with fish bones in its stomach. In addition, phytosaurs 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 Proterochampsia Middle Triassic reptiles of Asia Triassic China Fossils of China Fossil taxa described in 2016 Paleontology in Yunnan