| Pistosaurus | |
|---|---|
 | |
| Fossil | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Superorder: | †Sauropterygia |
| Family: | †Pistosauridae Zittel, 1887 |
| Genus: | †Pistosaurus Meyer,1839 |
| Type species | |
| †Pistosaurus longaevus Meyer, 1839 | |
Pistosaurus (exact etymology uncertain[1]) is an extinct genus of aquaticsauropterygianreptile closely related toplesiosaurs. Fossils have been found in France and Germany, and date to theMiddle Triassic. It contains asingle species,Pistosaurus longaevus. Pistosaurus is known as the oldest "subaquatic flying" reptile on earth.
The skull ofPistosaurus generally resembles that of other Triassicsauropterygians. However, there are several synapomorphies that makePistosaurus distinguished: the long, slender,snout; the possession of splint-like nasals that are excluded from the external naris; and the posterior extension of the premaxilla to the frontals.[2] Based on synapomorphies such as the small nasals size and the presence of interpterygoid vacuity,Pistosaurus is more closely related toPlesiosauria than toNothosaurus.[2]
Pistosaurus is often mistaken withNothosaurus and Plesiosauria.Nothosaurus belongs to the clade Nothosauroidea from the middle Triassic (approximately 199-251 million years ago); whilePistosaurus belongs to stem group Plesiosauria; and bothPistosaurus and Plesiosauria belongs to cladePistosauroidea from Triassic. Both Nothosauroidea and Pistosauroidea belong to Sauropterygia.[2]
Pistosaurus was about 3 metres (10 ft) long, and had a body form resembling that ofnothosaurs, aquatic reptiles that flourished during theTriassic. However, thevertebral column was stiff, like that of a plesiosaur, implying that the animal used its paddle-like flippers to propel itself through the water, as the plesiosaurs probably did. The head also resembled that of a plesiosaur, but with the primitivepalate of a nothosaur, and numerous, sharp teeth ideal for catching and eating fish.[3]
The description below is based on the specimen examined by paleontologist Sues in 1987.
The structure of pectoral girdle and humerus are used to support the anterior part of the body.[4] The scapula in pectoral girdle ofPistosaurus consists with a massive body and a short posterodorsal process. It is smaller in size compared to coracoid. And its lateral margin of the body is gently convex anteroposteriorly while the medial margin is more strongly convex.[4]
The coracoid bone ofPistosaurus is flat and expanded medially.[4] The glenoid region is similar toNothosaurus in development: both the slight notching of its margin and a distinct facet contact with the humeral head. There is also a ridge like thickening which links the glenoid to posteromedial region of the coracoid.[4] This feature is asynapomorphy that appears in plesiosaurs, which is a thickened ridge passes transversely across the anterior portion of the coracoid to connect the glenoid region. This feature is suggested related to compressional force by limb motion inPistosaurus.[4]
A specimen of the left humerus ofPistosaurus analyzed by Paleontologist A.R.I. Cruickshank is one of the largest specimens recorded: 245mm long and 45mm wide at the mid-shaft.[5] The specimen showing that the axis ofPistosaurus' humerus is straight, with the distal end slightly expanded posteriorly.[5] From proximal view, the head of the humerus is concave, which is a sign of a substantial cap of cartilage at the head of humerus. The humerus ofPistosaurus also lacksentepicondylar foramen.[5]
Pistosaurus has a strongly flattenedulna. It has medium length and nearly symmetrical in dorsal view.[4] Its anterior margin is more curved and thicker than the posterior one. This feature broads the wide spatium interosseum enclosed between radius and ulna.[4] The proximal end ofradius is less expanded than that of ulna, while the distal end is less expanded than proximal one but thickened.[4] The anterior margin is nearly straight while the posterior margin is more curved compared to the anterior one. Like other sauropterygians, the radius ofPistosaurus is slightly longer than the ulna.[4]
Thepelvic girdle ofPistosaurus is more similar to primitive sauropterygians than to plesiosaurs.
Theilium ofPistosaurus has an iliac blade, which has almost parallel anterior and posterior margins.[4] Same as other non-plesiosaur sauropterygians, the ilium inPistosaurus contacts both the pubis and the ischium, forming a ring-like structure. The ilium fromPistosaurus is relatively large in size compared toNothosaurus, whose ilia did not appear to have any elongated blade.[4]
The femur ofPistosaurus is longer than its humerus. Its anterior margin is almost straight whereas the posterior margin is concave.[4] According to the specimen provided by paleontologist Sues, the proximal articular end is much more robust than the distal one, and is more or less triangular in transverse section.[4]
Although it is unlikely thatPistosaurus was a direct ancestor of the plesiosaurs, the mixture of features suggests that it was closely related to that group.[3]
The followingcladogram follows an analysis by Ketchum & Benson, 2011.[6]
The classification forPlesiosauria was difficult at the first place. The anatomy of stem groupSauropterygia has very primitive synapomorphies such as dermal palate. Initially, Plesiosauria were suggested related toPistosauroidea, which belongs to Eusauropterygia from Triassic. Three genera of Plesiosauria was known in the history:Corosaurus alvocensis, Cymatosaurus, andPistosaurus longaevus.[7] A later discovery of a newPistosauridea from middle triassic of Nevada by paleontologist Sander indicates thatAugustasaurus is closely related toPistosaurus, while there are several difference including axial skeleton.[8]
| Pistosauria |
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There are several different ways for aquatic tetrapods to counteract their positive buoyancy caused by their lungs:pachyostosis,osteosclerosis,pachyosteosclerosis, and calcified cartilage of bone. The ultimate goal of these processes are to increase density for different parts of the body to offset the buoyancy, in order to live in an aquatic/semi-aquatic environment.[2] Bone histology ofPistosaurus longaevus studied by Paleontologist Krahl showed that the medullary region of humeri was filled, and it contained calcified cartilage incorporated into endoseal bone. According to Krahl, the small region of medullary of humeri is results from a suppressed perimedullary resorption activity, which is associated withosteosclerosis.[2]
Paleontologist Diedrich examined other pectoral and pelvic girdle ofPistosaurus. Together with the muscle grooves, they determined that a slight subaquatic flying starts withPistosaurus. And most of the propulsion occurs on the hindlimbs. The presence of enlargedcoracoid andpubis bone in pelvic girdle indicated that there were possibility forPistosaurus to develop flipper-like extremities.[9] What's more, the underwater flying mode suggested by paleontologist Michael. A. Taylor indicates that the left and right limb ofPlesiosaurus would simultaneously beat together. This feature contrasts to the terrestrial reptile who use right and left limbs for locomotion alternatively.[10]
The non-PlesiosaurianSauropterygians are found in various locations in China, Europe, America, Palestine and Tunisia. Although Nothosaurus, which is closely related toPistosaurus, are found plenty across Europe,Pistosaurus skull is only found in Germanic basin in Upper Muschelkalk.[2]
The early discovery ofPistosaurus skull was by H. v. Meyer. He discovered two skulls and a postcranial skeleton at the same location, possibly fromPistosaurus. Later after that, a new and well preserved postcranial skeleton was also found at the same location as previous specimens.[11] Paleontologist Geissler first described the skeletons and then paleontologist Strunz developed new hypothesis based on that. This skeleton was originally preserved at Strunz collection in Senckenberg Museum at Frankfurt a/M.[11]
AlthoughPistosauroidea has long been considered as structural antecedents ofPlesiosauria, a new specimen ofAugustasaurus discovered by paleontologist Sandra from Nevada had raised to against this theory. Opposed to previous hypothesis, the forelimb ofAugustasaurus was greatly reduced compared toPlesiosaurus. Therefore, Pistosauroidea was removed from stem group Plesiosauria and becomes paraphyletic group to Plesiosauria.[8]
Avascular necrosis, also known as bone necrosis, is associated with decompression syndrome (DCS). It is caused by expose rapid decrease of external pressure as well as rapid ascent in water column.[12] There features are often recognized in TriassicSauropterygians. According to paleontologist Surmik, the presence of decompression syndrome-related avascular necrosis inPistosaurus forelimb suggested thatPistosaurus used to live in aquatic or semi-aquatic environment. He also stated the possibility ofPistosaurus distributed in open marine cold water, and their effective metabolism is one of the reasons whyPistosaurus can survive the open sea.[12]
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