Soki Hattori (2016)
Evolution of the hallux in non-avian theropod dinosaurs.
Journal of Vertebrate Paleontology (advance online publication)
The emergence of a reversed hallux (first pedal digit) in the theropod lineage is regarded as an important indicator of increasing grasping ability and possibly arboreality. However, functions of the pes with a plesiomorphic, non-reversed hallux in non-avian theropods, other than the major role in supporting and propelling the body during the stance phase, have not been fully addressed. To clarify hallucal function, 21 specimens of non-avian theropods were examined in this study. Ancestrally in Theropoda, the proximal end of the first metatarsal reached the ankle joint as in extant crocodilians. The size of the first metatarsal was then reduced, lost direct contact with the ankle joint, and shifted more plantarly in basal neotheropods. In articulated coelophysoid specimens, the proximal end of the first metatarsal attaches to the medioplantar margin of the second metatarsal, whereas the distal part of the first metatarsal and phalanges of the hallux lies on the plantar aspect of the pes. In four articulated specimens of the dromaeosaurid Velociraptor, preserved positions of the first metatarsal on the second metatarsal vary from the medial to plantar aspects, and correspondingly the rotational axis of the extension/flexion movement at its distal articulation ranges from mediolateral to dorsoplantar. Similar conditions are also observed in troodontid specimens, suggesting the presence of intermetatarsal mobility between the first and second metatarsals enabled by a convex proximal articular surface of the first metatarsal in deinonychosaurians similar to the one present in extant birds.