Anatomy and relationships of the bizarre Early Cretaceous pliosaurid Luskhan itilensis

Pliosaurid plesiosaurians are iconic marine reptiles that regulated marine trophic chains from the Middle Jurassic to the early Late Cretaceous. However, their evolution during the Cretaceous remains poorly documented. Recent discoveries from the Hauterivian–Aptian interval suggest that the radiation of brachaucheniine pliosaurids produced a wide disparity of forms following the Pliosaurus-dominated assemblages of the Late Jurassic. Among the most bizarre of these early brachaucheniines is Luskhan itilensis, from the Hauterivian of Russia. We describe the osteology of this tusked, longirostrine pliosaurid and discuss its possible behaviour by drawing comparisons with other marine amniotes possessing forward-pointing teeth. We take this opportunity to make extensive anatomical comparisons among Cretaceous pliosaurids, including previously overlooked cranial features. Bayesian inference of phylogenetic relationships of plesiosaurians reveals that the internal branches in Late Jurassic–Late Cretaceous pliosaurids have generally low rates of morphological evolution, indicating that the recently described Early Cretaceous pliosaurids have effectively bisected the long branch leading to the ‘classical’ brachaucheniines of the middle Cretaceous (Brachauchenius, Kronosaurus and Megacephalosaurus). Pliosaurids exhibit low evolutionary rates and a dwindling disparity before their extinction, mirroring the events seen, roughly at the same time, for ichthyosaurians.