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''Dianopachysaurus'' 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
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 ...
known from the lower
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) of
Yunnan Province Yunnan , () is a landlocked province in 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 Kunming. The province borders the C ...
, 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 was found in the Middle
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
Lagerstatte 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 ...
. It was first named by Jun Liu, Olivier Rieppel, Da-Yong Jiang, Jonathan C. Aitchison, Ryosuke Motani, Qi-Yue Zhang, Chang-Yong Zhou and Yuan-Yuan Sun 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 is ''Dianopachysaurus dingi,'' thanking a Professor Ding for his help. ''Dianopachysaurus'' is most closely related to ''
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 Ch ...
'', another Chinese pachypleurosaur. Both belong to the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Keichousauridae. Pachypleurosaurs are hypothesized to have originated in the eastern
Tethys Ocean The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...
(South China) before spreading and diversifying in the western Tethys in what is now
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
. A large
ghost lineage A ghost lineage is a hypothesized ancestor in a species lineage that has left no fossil evidence yet can be inferred to exist because of gaps in the fossil record or genomic evidence. The process of determining a ghost lineage relies on fossilized ...
of eastern pachypleurosaurs has long been inferred based on the phylogeny of the group. ''Dianopachysaurus'' represents an early stage in the radiation of pachypleurosaurs and its early age fills in much of the gap.


Features

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 ...
, and only
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
so far, is an almost complete articulated skeleton. It measured long from nose tip to the last
caudal 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 i ...
known, although the end of the tail is missing after the 17th caudal vertebra. It is estimated to have measured up to in total body length and weighed .


Skull

The
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 ...
is broad and flat, with the
eyes Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conve ...
quite far forward and the snout shorter than the postorbital region. Much of the
occipital 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 cereb ...
region has been covered by the displaced
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
, so it is not visible and cannot be studied easily, but there appears to be no occipital crest. The
premaxillae 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 b ...
are quite broad, and form the anterior margin of the
naris 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 ...
. They show some weak signs of snout constriction, being slightly narrower than the rest of the skull, but it is not very pronounced. At the very anterior end of the
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * 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 ships * Ros ...
they are fused. The
maxillae 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. The t ...
are partially covered, and only a few maxillary
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
are at all visible, all smaller than the premaxillary teeth. They extend back, forming the posterolateral border of the naris and the lateral border of the
orbit In celestial mechanics, an orbit 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 object or position in space such as a p ...
to just beyond the midpoint. A few striations are visible on the maxillae. The nares are small, only long, and almost square or trapezoid. The
nasal bones The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
are elongated and thin, but still shorter than the frontals. They are separated anteriorly by the posterior process of the premaxillae, but meet in a suture posteriorly. There are many small pits present on their surfaces. The orbit is large, at in diameter, but has no scleral ossicles present. The prefrontals form the anterior border of the orbit, and are raised in a large ridge just in front of it. The frontals are fully fused into one large bone, and this forms the small ridge between the orbits, which were on the top of the head. Its most noticeable feature is the two processes extending backwards into the parietal almost up to the
pineal foramen A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhythm ...
. The postfrontals are triradiate and come between the orbits and the upper temporal fenestrae, but their anterior processes are mostly covered by the postorbitals. The jugals are very reduced, only thin slivers of bone bounding the orbits and just touching the upper temporal fenestrae. They form no part of the upper temporal arch. The upper temporal fenestrae are only 4.5 mm long, and much thinner, making them relatively small. There is only one parietal, as it is completely fused, and this bounds much of both upper temporal fenestrae and forms most of the upper temporal arch. It is quite broad and flat, but uniquely to this species there is a weak constriction where it meets the squamosals. The pineal foramen is close to the centre of the parietal, but slightly anteriorly displaced. Very small pits are present on the surface of the parietal bone. The postorbitals are quite small, and there is weak sculpturing on the posterior processes. The squamosals, which form the posterior margins of the upper temporal fenestrae, are large and triradiate, with forked anterior processes. Their lateral processes just descend to meet the
lower jaw In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
. The
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
is barely exposed at all, but the quadrate is just noticeable near the location of the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
. Only part of the
supraoccipital 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 cereb ...
is exposed, but it has a deeply concave posterior margin. The
surangular The suprangular or surangular is a jaw bone found in most land vertebrates, except mammals. Usually in the back of the jaw, on the upper edge, it is connected to all other jaw bones: dentary, angular, splenial and articular The articular bone i ...
, which is only exposed in the postorbital region, has no lateral ridge, and the angular is very small. A small fragment of
splenial The splenial is a small bone in the lower jaw of reptiles, amphibians and birds, usually located on the lingual side (closest to the tongue) between the angular and surangular The suprangular or surangular is a jaw bone found in most land verteb ...
is exposed, but the dentary is barely visible and no teeth can be seen.


Vertebrae

20 cervical vertebrae, 19
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
, 3
sacral vertebrae The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
and at least 17 caudal vertebrae are present. The
atlas An atlas is a collection of maps; it is typically a bundle of maps of Earth or of a region of Earth. Atlases have traditionally been bound into book form, but today many atlases are in multimedia formats. In addition to presenting geographic ...
,
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis *Axis (mathematics), a designator for a Cartesian-coordinate ...
and eighth-tenth cervical vertebrae are disarticulated, as is the posterior-most region of the tail (in fact missing). The dorsal vertebrae have no elongated
transverse processes 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 i ...
, but the zygapophyses are very pachyostotic.


Ribs

The cervical ribs are double-headed and start from the third cervical vertebra. The dorsal ribs are single-headed, significantly more elongate, and highly pachyostotic. The last dorsal rib is noticeably shorter than the preceding one, but still much longer than the following sacral ribs. It points laterally instead of trending towards the ilium. The sacral ribs are all articulated with the ilium but do not fuse to their vertebrae. The first caudal rib is far away from the ilium, but trends towards it.


Pectoral girdle and forelimbs

The
clavicles The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the right ...
are attached to the medial edge of the
scapulae 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 either ...
. The scapulae have highly developed dorsal wings, which are posteriorly directed and taper to blunt points. The proximal ends of the
humeri The humerus (; ) 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 a round ...
are mostly covered by the scapulae, and the deltapectoral flanges are not highly developed. The humeri are 10 mm long, and have unequal attachments for the
ulnae The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of the ...
and
radii In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
. The ulnae is no wider than the radii are, and they are both 5 mm long. An unknown carpal bone, probably an intermedium as it is quite elongate, is preserved between the ulna and radius of the right forelimb and there are three rounded carpal elements preserved in the left forelimb, one of which is the
ulnare The triquetral bone (; also called triquetrum, pyramidal, three-faced, and formerly cuneiform bone) is located in the wrist on the medial side of the proximal row of the carpus between the lunate and pisiform bones. It is on the ulnar side of the ...
. The
phalangeal formula The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
is not known due to incomplete preservation.


Pelvic girdle and hindlimbs

The ilia have a triangular
acetabulum The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint. Structure There are three bones of the ''os coxae'' (hip bone) that c ...
, and a reduced but still quite robust dorsal process. The pubis is round, and has an open obdurator foramen, but the
ischium The ischium () form ...
is almost entirely hidden. The
femurs The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with t ...
are slender and slightly
sigmoid Sigmoid means resembling the lower-case Greek letter sigma (uppercase Σ, lowercase σ, lowercase in word-final position ς) or the Latin letter S. Specific uses include: * Sigmoid function, a mathematical function * Sigmoid colon, part of the l ...
, with distinct striations on both ends. They are 14 mm long. The
fibulae The fibula 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. Its upper extremity is ...
are slightly longer than the
tibiae The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, at 6.5 rather than 6 mm long. The astragali are larger than the calcanea, but both have flat proximal margins. The first
metatarsals The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the medi ...
are distinctly shorter than the others, and many metatarsals are covered by phalanges and so not fully visible. Enough phalanges are preserved in their original positions, however, to let us see that all the most distal phalanges have pointed tips, which may originally have been claws, and to give a probable phalangeal formula for the foot of 2,3,4,5,4.


References

{{Taxonbar, from=Q3936027 Fossil taxa described in 2011 Pachypleurosaurs Triassic sauropterygians Middle Triassic reptiles of Asia Anisian life Guanling Formation Sauropterygian genera