A new pterosaur from the Middle Jurassic of Skye, Scotland and the early diversification of flying reptiles

The Middle Jurassic was a critical time in pterosaur evolution, witnessing the appearance of major morphological innovations that underpinned successive radiations by rhamphorhynchids, basally branching monofenestratans, and pterodactyloids. Frustratingly, this interval is particularly sparsely sampled, with a record consisting almost exclusively of isolated fragmentary remains. Here, we describe new material from the Bathonian-aged Kilmaluag Formation of Skye, Scotland, which helps close this gap. Ceoptera evansae (gen. et sp. nov.) is based on a three-dimensionally preserved partial skeleton, which represents one of the only associated Middle Jurassic pterosaurs. Ceoptera is among the first pterosaurs to be fully digitally prepared, and mu CT scanning reveals multiple elements of the skeleton that remain fully embedded within the matrix and otherwise inaccessible. It is diagnosed by unique features of the pectoral and pelvic girdle. The inclusion of this new Middle Jurassic pterosaur in a novel phylogenetic analysis of pterosaur interrelationships provides additional support for the existence of the controversial clade Darwinoptera, adding to our knowledge of pterosaur diversity and evolution.