Abstract
The feet are the only contact between the body and the substrate in limbed animals and as such they provide a crucial interface between the animal and its environment. This is especially true for bipedal and arboreal species living in a complex three-dimensional environment that likely induces strong selection on foot morphology. In birds, foot morphology is highly variable, with different orientations of the toes, making it a good model for the study of the role of functional, developmental, and phylogenetic constraints in the evolution of phenotypic diversity. Our data on the proportions of the phalanges analyzed in a phylogenetic context show that two different morphological patterns exist that depend mainly on habitat and toe orientation. In the anisodactyl foot, the hallux is the only backward-oriented toe and is enlarged in climbing species and reduced in terrestrial ones. Moreover, a proximo-distal gradient in phalanx size is observed depending on the degree of terrestriality. In the two other cases (heterodactyl and zygodactyl) that have two toes that point backward, the hallux is rather small in contrast to the other backward-pointing toe, which is enlarged. The first pattern is convergent and common among tetrapods and follows rules of skeletal development. The second pattern is unique for the clade and under muscle-morphogenetic control. In all cases, the functional result is the same tool, a pincer-like foot.