Deep-time invention and hydrodynamic convergences through amniote flipper evolution.

The diapsid plesiosaurs were pelagic and inhabited the oceans from the Triassic to the Cretaceous. A key evolutionary character of plesiosaurs is the four wing-like flippers. While it is mostly accepted that plesiosaurs were underwater fliers like marine turtles, penguins, and maybe whales, other swimming styles have been suggested in the past. These are rowing and a combination of rowing and underwater flight (e.g., pig-nosed turtle, sea lion). Underwater fliers use lift in contrast to rowers that employ drag. For efficiently profiting of lift during underwater flying, it is necessary that plesiosaurs twisted their flippers by muscular activity. To research the evolution of flipper twisting in plesiosaurs and functionally analogous taxa, including turtles, we used anatomical network analysis (AnNA) and reassessed distal flipper muscle functions. We coded bone-to-bone and additionally muscle-to-bone contacts in N × N matrices for foreflippers of the plesiosaur, the loggerhead sea turtle, the pig-nosed turtle, the African penguin, the California sea lion, and the humpback whale based on literature data. In "R," "igraph" was run by using a walktrap algorithm to obtain morphofunctional modules. AnNA revealed that muscle-to-bone contacts are needed to detect contributions of modules to flipper motions, whereas only-bone matrices are not informative for that. Furthermore, the plesiosaur, the marine turtles, the seal, and the penguin flipper twisting mechanisms, but the penguin cannot actively twist the flipper trailing edge. Finally, the foreflipper of the pig-nosed turtle and of the whale is not actively twisted during swimming.