Avemetatarsalia


Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians.[2] The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinosaurs were the largest terrestrial animals for much of the Mesozoic Era, and one group of small feathered dinosaurs (Aves, i.e. birds) has survived up to the present day. Pterosaurs were the first flying vertebrates and persisted through the Mesozoic before dying out at the Cretaceous-Paleogene (K-Pg) extinction event. Both dinosaurs and pterosaurs appeared in the Triassic Period, shortly after avemetatarsalians as a whole.[2][3] The name Avemetatarsalia was first established by British palaeontologist Michael Benton in 1999. An alternate name is Panaves, or "all birds", in reference to its definition containing all animals, living or extinct, which are more closely related to birds than to crocodilians.[4]

Although dinosaurs and pterosaurs were the only avemetatarsalians to survive past the end of the Triassic, other groups flourished during the Triassic. The most basal (earliest-branching) and plesiomorphic ("primitive") known avemetatarsalians were the aphanosaurs. Aphanosaurs were rare, four-legged carnivores which were only properly distinguished as a group in 2017.[5] The split between dinosaurs and pterosaurs occurred just after aphanosaurs branched off the archosaur family tree. This split corresponds to the subgroup Ornithodira, defined as the last common ancestor of dinosaurs and pterosaurs, and all of its descendants. Until the discovery of aphanosaurs, Ornithodira and Avemetatarsalia were considered roughly equivalent concepts.[3]

Pterosauromorpha includes all avemetatarsalians closer to pterosaurs than to dinosaurs. True non-pterosaur pterosauromorphs have been historically difficult to determine. Small, insectivorous archosaurs of the family Lagerpetidae may potentially be examples, alongside the similar genus Scleromochlus.[6] Dinosauromorpha, on the other hand, includes all avemetatarsalians closer to dinosaurs than to pterosaurs. Probable non-dinosaur dinosauromorphs include the diverse and widespread silesaurids, as well as more controversial and fragmentary taxa such as Marasuchus, Lagosuchus, Nyasasaurus, and Saltopus. Lagerpetids were also traditionally considered dinosauromorphs,[7][3] though this has been more recently debated.[8][6][9]

The foundational characteristic is the "advanced mesotarsal" ankles, which are characterized by a large astragalus and a small calcaneum. This ankle orientation operated on a single hinge, allowing for better mobility. Probably as a result of this change, the common ancestor of the avemetatarsalians had an upright, bipedal posture, with their legs extending vertically, similar to that of mammals.

Feathers and other filamentary structures are known across the avemetatarsalians, from the downy pycnofibers of pterosaurs, to quill-like structures present in ornithischian dinosaurs, such as Psittacosaurus and Tianyulong, to feathers in theropod dinosaurs and their descendants, birds.

Two clades of avemetatarsalians, pterosaurs and birds, independently evolved flight. Pterosaurs are the earliest vertebrates known to have evolved powered flight. Their wings are formed by a membrane of skin, muscle, and other tissues stretching from the ankles to a dramatically lengthened fourth finger.[10] Birds evolved flight much later. Their wings formed from elongated fingers and their arms, all covered with flight feathers.


The "advanced mesotarsal" ankle present in most avemetatarsalians.