The morphological basis of hallucal orientation in extant birds

The perching foot of living birds is commonly characterized by a reversed or opposable digit I (hallux). Primitively, the hallux of nonavian theropod dinosaurs was unreversed and lay parallel to digits II-IV. Among basal birds, a unique digital innovation evolved in which the hallux opposes digits II-IV. This digital configuration is critical for grasping and perching. I studied skeletons of modern birds with a range of hallucal designs, from unreversed (anteromedially directed) to fully reversed (posteriorly directed). Two primary correlates of hallucal orientation were revealed. First, the fossa into which metatarsal I articulates is oriented slightly more posteriorly on the tarsometatarsus, rotating the digit as a unit. Second, metatarsal I exhibits a distinctive torsion of its distal shaft relative to its proximal articulation with the tarsometatarsus, reorienting the distal condyles and phalanges of digit I. Herein, I present a method that facilitates the re-evaluation of hallucal orientation in fossil avians based on morphology alone. This method also avoids potential misinterpretations of hallucal orientation in fossil birds that could result from preserved appearance alone.