Pistosaurus

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Pistosaurus
Temporal range: Middle Triassic, Anisian–Ladinian
Pistosaurus longaevus Tubingen.JPG
Fossil
Scientific classification OOjs UI icon edit-ltr.svg
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 longaevus skull (cast) Pistosaurus skull.jpg
Pistosaurus longaevus skull (cast)

Pistosaurus (pistos in Greek meaning 'credible' and sauros 'lizard' [1] ) is an extinct genus of aquatic sauropterygian reptile closely related to plesiosaurs. Fossils have been found in France and Germany, and date to the Middle Triassic. It contains a single species, Pistosaurus longaevus. Pistosaurus is known as the oldest "subaquatic flying" reptile on earth.

Contents

The skull of Pistosaurus generally resembles that of other Triassic sauropterygians. However, there are several synapomorphies that make Pistosaurus 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 to Plesiosauria than to Nothosaurus . [2]

Pistosaurus is often mistaken with Nothosaurus and Plesiosauria. Nothosaurus belongs to the clade Nothosauroidea from the middle Triassic (approximately 199-251 million years ago); while Pistosaurus belongs to stem group Plesiosauria; and both Pistosaurus and Plesiosauria belongs to clade Pistosauroidea from Triassic. Both Nothosauroidea and Pistosauroidea belong to Sauropterygia. [2]

Description and paleobiology

Restoration of Pistosaurus longaevus Pistosaurus BW.jpg
Restoration of Pistosaurus longaevus

Pistosaurus was about 3 metres (10 ft) long, and had a body form resembling that of nothosaurs, aquatic reptiles that flourished during the Triassic. However, the vertebral 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 primitive palate of a nothosaur, and numerous, sharp teeth ideal for catching and eating fish. [3]

Post-cranial skeleton

The description below is based on the specimen examined by paleontologist Sues in 1987.

Pectoral girdle and forelimbs

The structure of pectoral girdle and humerus are used to support the anterior part of the body. [4] The scapula in pectoral girdle of Pistosaurus 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 of Pistosaurus is flat and expanded medially. [4] The glenoid region is similar to Nothosaurus 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 a synapomorphy 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 in Pistosaurus. [4]

A specimen of the left humerus of Pistosaurus 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 of Pistosaurus' 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 of Pistosaurus also lacks entepicondylar foramen . [5]

Pistosaurus has a strongly flattened ulna. 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 of radius 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 of Pistosaurus is slightly longer than the ulna. [4]

Pelvic girdle

The pelvic girdle of Pistosaurus is more similar to primitive sauropterygians than to plesiosaurs.

The ilium of Pistosaurus has an iliac blade, which has almost parallel anterior and posterior margins. [4] Same as other non-plesiosaur sauropterygians, the ilium in Pistosaurus contacts both the pubis and the ischium, forming a ring-like structure. The ilium from Pistosaurus is relatively large in size compared to Nothosaurus, whose ilia did not appear to have any elongated blade. [4]

The femur of Pistosaurus 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]

Classification

Limb bone Pistosaurus longaevus from Caldonazzo.JPG
Limb bone

Although it is unlikely that Pistosaurus was a direct ancestor of the plesiosaurs, the mixture of features suggests that it was closely related to that group. [3]

The following cladogram follows an analysis by Ketchum & Benson, 2011. [6]

The classification for Plesiosauria was difficult at the first place. The anatomy of stem group Sauropterygia has very primitive synapomorphies such as dermal palate. Initially, Plesiosauria were suggested related to Pistosauroidea , which belongs to Eusauropterygia from Triassic. Three genera of Plesiosauria was known in the history: Corosaurus alvocensis, Cymatosaurus, and Pistosaurus longaevus. [7] A later discovery of a new Pistosauridea from middle triassic of Nevada by paleontologist Sander indicates that Augustasaurus is closely related to Pistosaurus, while there are several difference including axial skeleton. [8]

  Pistosauria  

"Pistosaurus postcranium"

Augustasaurus hagdorni

Bobosaurus forojuliensis

Pistosaurus

Yunguisaurus liae

 Plesiosauria 
  Pliosauroidea  
  Pliosauridae  

Thalassiodracon hawkinsii

Hauffiosaurus spp.

Attenborosaurus conybeari

advanced pliosaurids

  Rhomaleosauridae  

Anningasaura lymense

advanced rhomaleosaurids

"Plesiosaurus" macrocephalus

Archaeonectrus rostratus

Macroplata tenuiceps

  Plesiosauroidea  

Stratesaurus taylori

  Plesiosauridae  

Seeleyosaurus guilelmiimperatoris

OUMNH J.28585

Plesiosaurus dolichodeirus

Elasmosauridae and Cryptoclidia

Microcleidus homalospondylus

Hydrorion brachypterygius

Occitanosaurus tournemiensis

Geological environment information

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 of Pistosaurus 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 with osteosclerosis. [2]

Paleontologist Diedrich examined other pectoral and pelvic girdle of Pistosaurus. Together with the muscle grooves, they determined that a slight subaquatic flying starts with Pistosaurus. And most of the propulsion occurs on the hindlimbs. The presence of enlarged coracoid and pubis bone in pelvic girdle indicated that there were possibility for Pistosaurus 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 of Plesiosaurus would simultaneously beat together. This feature contrasts to the terrestrial reptile who use right and left limbs for locomotion alternatively. [10]

Historical information and discovery

The non-Plesiosaurian Sauropterygians are found in various locations in China, Europe, America, Palestine and Tunisia. Although Nothosaurus, which is closely related to Pistosaurus, are found plenty across Europe, Pistosaurus skull is only found in Germanic basin in Upper Muschelkalk. [2]

The early discovery of Pistosaurus skull was by H. v. Meyer. He discovered two skulls and a postcranial skeleton at the same location, possibly from Pistosaurus. 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]

Although Pistosauroidea has long been considered as structural antecedents of Plesiosauria, a new specimen of Augustasaurus discovered by paleontologist Sandra from Nevada had raised to against this theory. Opposed to previous hypothesis, the forelimb of Augustasaurus was greatly reduced compared to Plesiosaurus . 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 Triassic Sauropterygians. According to paleontologist Surmik, the presence of decompression syndrome-related avascular necrosis in Pistosaurus forelimb suggested that Pistosaurus used to live in aquatic or semi-aquatic environment. He also stated the possibility of Pistosaurus distributed in open marine cold water, and their effective metabolism is one of the reasons why Pistosaurus can survive the open sea. [12]

Related Research Articles

<i>Plesiosaurus</i> Extinct genus of reptiles

Plesiosaurus is a genus of extinct, large marine sauropterygian reptile that lived during the Early Jurassic. It is known by nearly complete skeletons from the Lias of England. It is distinguishable by its small head, long and slender neck, broad turtle-like body, a short tail, and two pairs of large, elongated paddles. It lends its name to the order Plesiosauria, of which it is an early, but fairly typical member. It contains only one species, the type, Plesiosaurus dolichodeirus. Other species once assigned to this genus, including P. brachypterygius, P. guilielmiimperatoris, and P. tournemirensis have been reassigned to new genera, such as Hydrorion, Seeleyosaurus and Occitanosaurus.

<span class="mw-page-title-main">Plesiosaur</span> Order of reptiles (fossil)

The Plesiosauria or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia.

<span class="mw-page-title-main">Sauropterygia</span> Group of Mesozoic aquatic reptiles

Sauropterygia is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria became extinct at the end of that period. The plesiosaurs would continue to diversify until the end of the Mesozoic. Sauropterygians are united by a radical adaptation of their pectoral girdle, adapted to support powerful flipper strokes. Some later sauropterygians, such as the pliosaurs, developed a similar mechanism in their pelvis.

<i>Nothosaurus</i> Extinct genus of reptiles

Nothosaurus is an extinct genus of sauropterygian reptile from the Triassic period, approximately 240–210 million years ago, with fossils being distributed from North Africa and Europe to China. It is the best known member of the nothosaur order.

<span class="mw-page-title-main">Shoulder girdle</span> Set of bones which connects the arm to the axial skeleton on each side

The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of the clavicle, scapula, and coracoid. Some mammalian species have only the scapula.

<i>Huabeisaurus</i> Extinct genus of dinosaurs

Huabeisaurus was a genus of dinosaur from the Late Cretaceous. It was a sauropod which lived in what is present-day northern China. The type species, Huabeisaurus allocotus, was first described by Pang Qiqing and Cheng Zhengwu in 2000. Huabeisaurus is known from numerous remains found in the 1990s, which include teeth, partial limbs and vertebrae. Due to its relative completeness, Huabeisaurus represents a significant taxon for understanding sauropod evolution in Asia. Huabeisaurus comes from Kangdailiang and Houyu, Zhaojiagou Town, Tianzhen County, Shanxi province, China. The holotype was found in the unnamed upper member of the Huiquanpu Formation, which is Late Cretaceous (?Cenomanian–?Campanian) in age based on ostracods, charophytes, and fission-track dating.

<i>Keichousaurus</i> Extinct genus of reptiles

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<i>Aristonectes</i> Extinct genus of marines reptiles

Aristonectes is an extinct genus of large elasmosaurid plesiosaurs that lived during the Maastrichtian stage of the Late Cretaceous. Two species are known, A. parvidens and A. quiriquinensis, whose fossil remains were discovered in what are now Patagonia and Antarctica. Throughout the 20th century, Aristonectes was a difficult animal for scientists to analyze due to poor fossil preparation, its relationships to other genera were uncertain. After subsequent revisions and discoveries carried out from the beginning of the 21st century, Aristonectes is now recognised as the type genus of the subfamily Aristonectinae, a lineage of elasmosaurids characterized by an enlarged skull and a reduced length of the neck.

<i>Augustasaurus</i> Extinct genus of reptiles

Augustasaurus is a genus of aquatic sauropterygian reptile belonging to the Pistosauria, a clade containing plesiosaurs and their close relatives. Pistosaurus and Augustasaurus were thought to be the only known members of the family Pistosauridae. However, some recent cladistic analyses found Augustasaurus to be a more advanced pistosaur, as a sister group of the order Plesiosauria. The only known species of Augustasaurus is Augustasaurus hagdorni, which was first described in 1997.

Bobosaurus is an extinct genus of sauropterygian reptile related to plesiosaurs. It is based on the holotype MFSN 27285, a partial skeleton found in Early Carnian-age rocks of the Rio del Lago Formation, northeastern Italy. Bobosaurus was named in 2006 by Fabio M. Dalla Vecchia and the type species is B. forojuliensis. It may be a pistosaurid, or closer to Plesiosauria. A recent cladistic analysis found it to be a pistosaur. It was relatively large animal, with more than 3 m (9.8 ft) in length.

<i>Tatenectes</i> Genus of extinct plesiosaur from the upper Jurassic

Tatenectes is a genus of cryptoclidid plesiosaur known from the Upper Jurassic of Wyoming. Its remains were recovered from the Redwater Shale Member of the Sundance Formation, and initially described as a new species of Cimoliosaurus by Wilbur Clinton Knight in 1900. It was reassigned to Tricleidus by Maurice G. Mehl in 1912 before being given its own genus by O'Keefe and Wahl in 2003. Tatenectes laramiensis is the type and only species of Tatenectes. While the original specimen was lost, subsequent discoveries have revealed that Tatenectes was a very unusual plesiosaur. Its torso had a flattened, boxy cross-section and its gastralia exhibit pachyostosis (thickening). The total length of Tatenectes has been estimated at 2–3 meters (6.6–9.8 ft).

<i>Plesiopterys</i> Extinct genus of reptiles

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<i>Simosaurus</i> Extinct genus of reptiles

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<span class="mw-page-title-main">Pistosauroidea</span> Extinct clade of reptiles

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<span class="mw-page-title-main">Microcleididae</span> Extinct family of reptiles

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<i>Atychodracon</i> Extinct genus of reptiles

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References

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