Thalattosauria

Thalattosaurs have moderate adaptations to marine lifestyles, including long, paddle-like tails and slender bodies with more than 20 dorsal vertebrae. There are few unique traits of the postcranial skeleton shared by all thalattosaurs, but the skeleton is still useful for distinguishing between askeptosauroids and thalattosauroids. Askeptosauroids are characterized by elongated necks with short neural spines and at least 11 vertebrae, while thalattosauroids have shorter necks sometimes involving as few as four vertebrae. Thalattosauroids also have tall neural spines on their neck, back, and especially the tail vertebrae, increasing the surface area for swimming via lateral undulation. Thalattosauroids additionally possess short, wide limb bones poorly adapted for movement on land. In this superfamily, the humerus is widest near the shoulder, the femur is widest near the knee, the radius is reniform ("kidney-shaped"), and phalanges are long and plate-like. Askeptosauroids retain hourglass-shaped limb bones like land reptiles, but even they share specializations with thalattosauroids such as a short tibia and fibula, with the latter expanding near the ankle. Thalattosaurs have a rostrum (snout) significantly longer than the portion of the skull behind the eyes. A majority of this length is formed from the premaxillary bones, and the nares (nostril holes) are shifted back close to the eyes. The premaxillae stretch back very far and are incised into the frontal bones. This leads to an unusual trait that is characteristic of thalattosaurs, where the left and right nasal bones are separated from each other and restricted to a small portion of the snout near the nares. The lacrimal bone is typically lost or fused to the large crescent-shaped prefrontal bone in front of the orbit, mirroring the postfrontal bone which is usually fused to the three-pronged postorbital bone behind the orbit. The exception to this rule is Gunakadeit, which has a straight snout and many slender teeth. Thalattosaurs often have a pronounced retroarticular process at the rear of the mandible. Thalattosauroids are more specialized than askeptosauroids in jaw anatomy, as they have evolved a large peak-like coronoid bone and an angular bone that extends far forwards along the lower edge of the jaw. Palatal dentition is extensive in thalattosauroids but absent in askeptosauroids. ==Paleobiology==

Thalattosaurs are only known from marine deposits, indicating that they were all primarily aquatic reptiles. The retracted nostrils and long, paddle-shaped tail are further evidence for aquatic habits. Thalattosauroids seemingly spent all of their time in the water, with short, wide limbs, poorly developed wrist and ankle bones, and tall vertebrae adapted for swimming via lateral undulation. Even so, they retained strong claws and functional digits which had not transformed into flippers, in contrast to ichthyosaurs and sauropterygians. Unlike these other marine reptiles, there is no evidence that thalattosaurs fully adapted to a pelagic life out in the open ocean, and instead they probably all lived in warm waters close to the coast. Askeptosauroids had stronger limbs more typical of terrestrial reptiles, indicating they would have been capable of moving around on land to some extent. They likely primarily used their tails when swimming, while thalattosauroids may have utilized their body and tail in conjunction. Thalattosaurs had diverse diets, though they probably all involved marine animals in one way or another. Endennasaurus probably predated small animals like fish fry or small crustaceans due to its lack of teeth. technique of correlating diet with tooth shape, the taller teeth were suited for a "crunching" diet, involving armored fish, large crustaceans, and thin-shelled ammonites. The low, robust teeth would have been useful for a "crushing" diet specialized in large molluscs or other thick-shelled prey. Gunakadeit's slender teeth correlated with the "Pierce II" guild of Massare (1987), indicating it likely fed on soft, fast-moving fish and squid. It also had a large hyoid apparatus which may have played a role in suction feeding. ==Distribution==

It is not certain where thalattosaurs originated from. During the Triassic period, the earth had one giant supercontinent, Pangaea, which was surrounded by the superocean Panthalassa. The eastern portion of Pangaea was incised by a massive tropical inland sea, the Tethys Ocean, which extended all the way from China to Western Europe. While thalattosauroids are known from worldwide Triassic marine deposits, askeptosauroids are only known in Tethyan deposits. Assuming Endennasaurus and Askeptosaurus were the most basal askeptosauroids, Askeptosauroidea would have originated in the Western Tethys Ocean, now the Alpine region of Europe. Trans-Panthalassa connections are also observed in other Triassic marine life such as pistosaurs and ammonites. Evidently thalattosaurs were capable of dispersing throughout major marine regions multiple times before the group's extinction, with thalattosauroids likely more prolific at spreading than askeptosauroids due to their greater aquatic adaptations. ==Classification==

Early hypotheses (such as lizards) or rhynchocephalians (such as the tuatara, pictured) When first named by Merriam in 1904, Thalattosauria was only known by the species Thalattosaurus alexandrae. Based primarily on the overall skull shape, it was hypothesized to have been close to the reptile order Rhynchocephalia, which includes Sphenodon (the living tuatara). Nevertheless, Thalattosaurus was recognized as distinct enough to be given its own order, and was tentatively grouped along with Rhynchocephalia in the group Diaptosauria, a collection of various "primitive" reptiles now known to be polyphyletic. Within Diaptosauria, thalattosaurs were also considered very closely related to choristoderes and "Proganosauria" (parareptiles). Comparisons were also made with Parasuchia (phytosaurs), Lacertilia (lizards), and Proterosuchus, but dismissed as incompatible with proposed evolutionary schemes. Further discussion by Merriam (1905) considered a relationship with ichthyosaurs due to their similar ecology, but questioned why their skull and vertebral anatomy would diverge so widely if they had a close common ancestor. He proposed that potential similarities were best explained as convergent evolution. The possibility that thalattosaurs diverged from reptiles close to lizards (such as Paliguana) was described in more detail, with thalattosaurs serving as a short-lived early attempt for near-lizards to return to the sea, an evolutionary process later repeated more successfully when mosasaurs evolved from true lizards. Nevertheless, Merriam found no clear evidence that any previously known reptile group was directly ancestral to thalattosaurs or vice versa. They were probably descended from land-dwelling Permian reptiles, and not closely related to other marine reptile groups which first evolved in the Triassic. Modern classification and external relationships , An analysis by Müller (2004) has even considered thalattosaurs to belong just outside of Sauria. Unusually, thalattosaurs have an affinity to shift near ichthyosaurs (in the group Ichthyosauromorpha) when certain basal saurians or near-saurians are excluded from the data set. Some analyses derived from Müller (2004) group thalattosaurs in a "marine superclade" with ichthyosauromorphs and sauropterygians, and sometimes with turtles, archosauromorphs, or lepidosauromorphs as well. For example, Simões et al (2022) classify thalattosaurs as the sister group of the sauropterygians, with their clade being sister to the ichthyosauromorphs, and all three being basal archosauromorphs. However, cladograms generated by these analyses change in unpredictable ways through alterations to their methodology (such as including or excluding aquatic adaptations or switching between parsimony and bayesian inference), leading some to have concerns over the validity of the "marine superclade". While thalattosaurs are almost certainly diapsids, the large degree of uncertainty surrounding their outgroup relations has led most modern paleontologists to classify them as Diapsida incertae sedis. Internal relationships One of the first phylogenetic analyses specifically focusing on thalattosaurs was part of Nicholls (1999)'s reevaluation of Thalattosaurus and Nectosaurus. She used a restricted definition of Thalattosauria which referred to a clade including all reptiles more closely related to Nectosaurus and Hescheleria than to Endennasaurus or Askeptosaurus. The more inclusive group including Askeptosaurus, Endennasaurus, and traditional thalattosaurs was given the name Thalattosauriformes.). Meanwhile, the clade containing reptiles closer to askeptosaurids is termed Askeptosauroidea'' However, uncertainty over Endennasaurus's thalattosaurian ancestry led to it being excluded from these analyses. After Müller et al. (2005) re-affirmed that Endennasaurus was closely related to Askeptosaurus, all thalattosaurs known at the time were finally combined into phylogenetic analyses. Studies by Rieppel, Liu, Cheng, Wu, and others continued to identify new Chinese taxa such as Miodentosaurus and various species of Anshunsaurus and Xinpusaurus, though homoplasy in these new taxa has led to little resolution in the structure of the two major branches of Thalattosauria. In an attempt to remedy this problem, new phylogenetic analyses were developed by Liu et al. (2013) during the description of Concavispina'', The internal relationships of thalattosaurs is still considered tentative and inconclusive, although the fundamental structure of the group (a monophyletic Thalattosauria clade split into askeptosauroids and thalattosauroids) is very stable. Some paleontologists have attempted to divide thalattosaurs into families. One family, Askeptosauridae, is typically considered to include Askeptosaurus and Anshunsaurus, with a few studies also placing Miodentosaurus or Endennasaurus within it. Another family, Thalattosauridae, was originally used to group Thalattosaurus and Nectosaurus, was later redefined to exclude Nectosaurus, and later still encompassed practically all thalattosauroids. Many thalattosaur-focused paleontologists avoid using family names due to their inconsistent usage and questionable validity. The following cladogram represents the results of a thalattosaur ingroup phylogenetic analysis by Druckenmiller et al. (2020). }} ==List of genera==

Other thalattosaurs include unnamed or indeterminate species from the Kössen Formation of Austria, and Pardonet Formations of British Columbia, the Natchez Pass Formation of Nevada, and the Vester Formation of Oregon. Blezingeria, a fragmentary marine reptile from the Muschelkalk of Germany, has also been considered a thalattosaur by some authors but this assignment is uncertain at best. Thalattosaurian fragments are known from Spanish Muschelkalk as well. Neosinasaurus, a poorly-known reptile from the Xiaowa Formation of China, has been identified as a possible thalattosaur. was described as Gunakadeit in 2020. ==References==