Qianosuchus Mixtus Skeletal
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''Qianosuchus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of aquatic
poposauroid Poposauroidea is a clade of advanced pseudosuchians. It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as '' Qianosuchus'' and ''Lotosaurus''. It excludes most large predatory quadrupedal "rauisuchian ...
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 ...
from the middle
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
(
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
)
Guanling Formation The Guanling Formation is a Middle Triassic (Anisian or Pelsonian in the regional chronostratigraphy) geologic Formation (geology), formation in southwestern China. Geology The formation encompasses two members. The first member is primarily c ...
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 Panzhou is divided int ...
,
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 ...
. It is represented by two nearly complete skeletons and a crushed skull preserved in the limestone. ''Qianosuchus'' was at least 3 metres long, and had several skeletal adaptations which indicate a semi-marine lifestyle, similar to modern-day
saltwater crocodile The saltwater crocodile (''Crocodylus porosus'') is a crocodilian native to saltwater habitats, brackish wetlands and freshwater rivers from India's east coast across Southeast Asia and the Sundaland to northern Australia and Micronesia. It ha ...
s. These adaptations have not been seen in any other archosaur from the Triassic.


Description

''Qianosuchus'' had a skull around 33 cm (13 inches) long, with an elongated snout. The
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit ** podium * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ...
formed by 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 ...
is shallow at the front of the skull but deepens posteriorly. Each premaxilla has nine long teeth, and 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 ...
e bear eighteen teeth each. All the teeth are laterally compressed, curved backwards and serrated, like those of most other carnivorous archosaurs. The nares are expanded and elongated and almost collide with 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, meaning that the septum (bony wall) between them is thin and lightweight. Unusually, the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
forms no part of the border of the antorbital fenestra. Each
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
had a large and well-developed
sclerotic ring The scleral ring or sclerotic ring is a hardened ring of plates, often derived from bone, that is found in the eyes of many animals in several groups of vertebrates. Some species of mammals, amphibians, and crocodilians lack scleral rings. The rin ...
in it, which would have reinforced the eyeball under pressure when ''Qianosuchus'' was diving. The
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
s have deep fossae (depressions) on their upper surface, which stretch backwards to the sutures with the parietals. Another such fossa is present between the two parietals themselves. The
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
turns down very slightly at the tip; a precise tooth count is unknown due to the bone being hidden by the maxillary teeth in the fossils. The
hyoid bone The hyoid-bone (lingual-bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid-cartilage. At rest, it lies between the base of the mandible and the third cervical verte ...
s are long and slender, with slightly expanded ends. ''Qianosuchus'' had nine cervical, fifteen
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
, two sacral and at least 50
caudal vertebrae Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx. In many reptiles, some of the caud ...
. The posterior end of the tail is missing in both skeletons. The
neural spines Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
grow taller posteriorly (further down the tail), making the caudal vertebrae tall but thin in that area. The first 23 caudal vertebrae have
transverse processes Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
, but these processes are lost further back. The vertebral centra grow shorter posteriorly, making the tail more flexible than the neck. Some of the more anterior caudal vertebrae have chevron bones ventral to them which also increase the height of the tail. All presacral vertebrae have 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 amph ...
s at the top of their neural spines. The
cervical rib Cervical ribs are the ribs of the neck in many tetrapods. In most mammals, including humans, cervical ribs are not normally present as separate structures. They can, however, occur as a pathology. In humans, pathological cervical ribs are usually no ...
s are elongate, at least four times the length of their corresponding centra, and may have had strong muscles attached enabling it to create suction in its throat while lunging forward at prey by expanding the
oesophagus The esophagus (American English), oesophagus (British English), or œsophagus ( archaic spelling) ( see spelling difference) all ; : ((o)e)(œ)sophagi or ((o)e)(œ)sophaguses), colloquially known also as the food pipe, food tube, or gullet, ...
. The dorsal ribs are expanded and pachyostotic at their distal ends. ''Qianosuchus
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 ...
e were thin and short, but had an extremely broad dorsal blade. Its coracoid bones were oval-shaped and quite thin, while its
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
s articulated almost at right angles with the
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals ( marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In ...
to form an L-shaped outline from the side. The
humeri The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of ...
were slender and lightly built, and almost totally straight. Neither partial skeleton has preserved the forelimbs below the elbows. The
pelvic girdle The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the Ilium (bone) ...
is similar to that of closely related but more terrestrial archosaurs, with the large posterior process and small anterior process on the ilium. The pubis had a deep foramen close to the proximal end, while the distal end of the thinner and shorter
ischium The ischium (; : is ...
was slightly expanded. The
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
was weakly sigmoid, and the
fibula The fibula (: fibulae or fibulas) 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. ...
and
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
were almost exactly the same length. 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 ...
had a hemicylindrical condyle and a broad calcaneal tuber, while 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 ...
had a convex facet for the tibia. Five
metatarsals The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges (toes). Lacking individual names, the metatarsal bones are nu ...
and two tarsals are known, with the fifth metatarsal slightly hooked.


Paleobiology

''Qianosuchus'' was well adapted to a semi-marine lifestyle, with a laterally compressed tail and tall neural spines providing a greater surface area, indicating an animal reliant on its undulating tail for propulsion. Its tail is actually more expanded than those of several other marine reptiles such as '' Hupehsuchus'' and the modern
marine iguana The marine iguana (''Amblyrhynchus cristatus''), also known as the sea iguana, saltwater iguana, or Galápagos marine iguana, is a species of Iguanidae, iguana found only on the Galápagos Islands (Ecuador). Unique among modern lizards, it is a m ...
, so ''Qianosuchus'' was almost certainly a competent swimmer. The thin scapulae and coracoids are also seen in many marine reptiles such as
ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s and
mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains were discovered in ...
, while the long neck and reduced dermal armour are seen in marine reptiles such as ''
Tanystropheus ''Tanystropheus'' (~ 'long' + 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic period, Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than t ...
.'' However, its pelvic girdle and large, relatively unspecialized legs would have allowed ''Qianosuchus'' to walk around on land as well, and may well have had an erect or semi-erect posture, based on the ankle joint. All this suggests that ''Qianosuchus'' lived a semi-aquatic lifestyle in and around the shallow seas where it lived, hunting either on water or on land.


References


External links


Hmnh.org: Triassic archosaur Qianosuchus was an ancient mariner
{{Taxonbar, from=Q593059 Poposauroidea Triassic archosaurs Middle Triassic reptiles of Asia Anisian life Fossil taxa described in 2006 Anisian genus first appearances Guanling Formation Prehistoric pseudosuchian genera