The locomotory apparatus and paraxial swimming in fossil and living marine reptiles: comparing Nothosauroidea, Plesiosauria, and Chelonioidea

Abstract

The terrestrial origins of the diapsid Sauropterygia and Testudines are uncertain, with the latter being highly controversially discussed to this day. For only 15 Ma, Nothosauroidea lived in shallow-marine seas of the Triassic. Contrastingly, the pelagic Plesiosauria evolved in the Late Triassic, dispersed globally, and inhabited the oceans of the Jurassic and Cretaceous for approximately 135 Ma. Since the Cretaceous (~ 100 Ma), Chelonioidea, the modern sea turtles, have populated the oceans. All three groups evolved aquatic paraxial locomotion. Nothosaurs swam with their foreflippers, supported by the swimming tail. Plesiosaurs are the only tetrapods to have ever evolved four hydrofoil-like flippers. The plesiosaur flipper beat cycle has been debated for nearly two centuries. The different proposed locomotory styles (rowing, rowing-flight, underwater flight) are discussed in this review. A fourth gait that is employed by Carettochelys insculpta, which combines rowing and flying, is introduced. The osteology of the locomotory apparatus of nothosaurs and plesiosaurs is reviewed and compared to that of extant underwater-flying Chelonioidea. In conclusion, underwater flight remains the favoured locomotory style for plesiosaurs. Also, the review reveals that nothosaur locomotion has largely remained unstudied. Further, our understanding of joint morphologies and mobilities of the foreflipper in nothosaurs, plesiosaurs, and even recent sea turtles, and of the hindflipper in plesiosaurs, is very limited. It is crucial to the discussion of locomotion, to find out, if certain limb cycles were even possible, as evidence seems to point to the improbability of a rowing motion because of limited humerus and femur long axis rotation in plesiosaurs.