A new anhanguerid pterosaur (Pterosauria, Pterodactyloidea) from the Lower Cretaceous Crato Formation of Brazil and its implications for Anhanguerinae radiation

Anhangueridae is one of the most diverse clades of pterodactyloid pterosaurs, with an outstanding fossil record in the Lower Cretaceous of the Araripe Basin, northeastern Brazil. However, diagnostic anhanguerid material from the Crato Formation remains scarce, limiting interpretations of the clade evolution within the Santana Group. Here, we describe Anhykenhe smaugi gen. et sp. nov., a new anhanguerine pterosaur from the Aptian Crato Formation, based on an almost complete skull associated with a lower jaw (CCNH 801). The new taxon is diagnosed by a distinctive combination of characters, some of which are previously unknown within Ornithocheiriformes. Phylogenetic analysis recovers Anhykenhe smaugi gen. et sp. nov. as the sister taxon to the Romualdo Formation anhanguerines. This result indicates that substantial diversification within Santana Group anhanguerines had already occurred by the late Aptian, with the Crato Formation preserving at least two distinct lineages, and suggests that the taxonomically rich anhanguerine assemblage of the Romualdo Formation reflects a subsequent expansion of this diversity.