Luskhan

With these features, it is the pliosaur that approaches closest to the distantly-related piscivorous polycotylids, having convergently evolved these traits more than 10 million years apart.

A nearly complete fossil skeleton of a pliosaur, preserved in three dimensions, was found by Gleb N. Uspensky in 2002 on the eastern bank of the Volga River, 3 kilometres (1.9 mi) north of the village of Slantsevy Rudnik in the Ulyanovsk region of western Russia.

Other authors were Valentin Fischer, Roger Benson, Nikolay Zverkov, Laura Soul, Maxim Arkhangelsky, Olivier Lambert, Ilya Stenshin, and Patrick Druckenmiller.

The genus name, Luskhan, is derived from luuses, spirits and masters of water in Mongolian and Turkic mythology, plus the suffix khan, meaning "chief".

Additional characteristics typical of brachauchenines include the parietal bone extending forward to the position of the nostrils; the snout being unconstricted, but bearing an expansion on the bottom surface like Megacephalosaurus; the retroarticular process at the back of the lower jaw being inturned; the teeth in the upper jaw being equally-sized; the facets that articulate with the cervical ribs on the cervical vertebrae being placed relatively high, similar to Kronosaurus and Brachauchenius but unlike Jurassic pliosaurs and other plesiosaurs; the transverse processes attaching to the dorsal vertebrae above the level of the neural canals; and the long coracoid measuring 2.3 times the length of the scapula (2.5 in Kronosaurus).

[2][8] Unusually, Luskhan also lacks many of the adaptations for hunting large prey seen in other brachauchenines: the snout is very thin; there is no keel on the bottom of the fused symphysis of the lower jaw; there is no diastema (or gap in the tooth row); the bones of the upper jaw (the premaxilla and maxilla) are expanded outwards; and there are no caniniform ("canine-like") teeth.

Unlike other thalassophoneans, but like the Elasmosauridae,[11] the blade of the scapula is relatively short, being only as tall vertically as the longitudinal distance from its base to the articulation with the coracoid.

[2] The following cladogram follows an analysis by Fischer and colleagues in 2017, based on a dataset published by Benson and Druckenmiller in 2014[3] that was previously modified for the description of Makhaira in 2015.

[2] Stratesaurus Rhomaleosauridae Plesiosauroidea Thalassiodracon Hauffiosaurus Attenborosaurus Anguanax Marmornectes Peloneustes "Pliosaurus" andrewsi Simolestes Liopleurodon Pliosaurus Makhaira Luskhan Stenorhynchosaurus Kronosaurus Megacephalosaurus Brachauchenius Along with Makhaira and Stenorhynchosaurus, Luskhan forms an evolutionary grade that fills a critical gap of 40 million years - from the Berriasian to the Barremian epochs of the Early Cretaceous - in the evolution of brachauchenines.

[12] Thus, Luskhan departed from the typical apex predator niche of thalassophoneans in order to colonize a lower trophic level, having done so independently 10 million years after polycotylids.

[2] Magnetostratigraphy indicates that this zone dates to the Hauterivian epoch of the early Cretaceous Period, approximately 128 million years ago.

[19] In terms of invertebrates, the S. versicolor zone is additionally characterized by the ammonites S. coronatiforme, S. pavlovae, S. intermedium, and S. polivnense; the bivalves Inoceramus aucella, Prochinnites substuderi, Astarte porrecta, and Thracia creditica; and the belemnites Acroteuthis pseudopanderi, Praeoxyteuthis jasikofiana, Aulacoteuthis absolutiformis, and A.

[1][20] The gastropods Ampullina sp., Avellana hauteriviensis, Claviscala antiqua, Cretadmete neglecta, Eucyclus sp., Hudlestonella pusilla, Khetella glasunovae, Sulcoactaeon sp., Tornatellaea kabanovi, Trilemma russiense, and Turbinopsis multicostulata have also been found within the S. versicolor zone around Ulyanovsk.

Holotype skull
Size comparison of the holotype