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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.

A common problem in the quantification of the orientation of the femoral neck is the difficulty to determine its true axis; however, this axis is typically estimated visually only. Moreover, the orientation of the femoral neck is commonly analysed using angles that are dependent on anatomical planes of reference and only quantify the orientation in two dimensions. The purpose of this study is to establish a method to determine the three-dimensional orientation of the femoral neck using a three-dimensional model. An accurate determination of the femoral neck axis requires a reconsideration of the complex architecture of the proximal femur. The morphology of the femoral neck results from both the medial and arcuate trabecular systems, and the asymmetry of the cortical bone. Given these considerations, two alternative models, in addition to the cylindrical one frequently assumed, were tested. The surface geometry of the femoral neck was subsequently used to fit one cylinder, two cylinders and successive cross-sectional ellipses. The model based on successive ellipses provided a significantly smaller average deviation than the two other models (P < 0.001) and reduced the observer-induced measurement error. Comparisons with traditional measurements and analyses on a sample of 91 femora were also performed to assess the validity of the model based on successive ellipses. This study provides a semi-automatic and accurate method for the determination of the functional three-dimensional femoral neck orientation avoiding the use of a reference plane. This innovative method has important implications for future studies that aim to document and understand the change in the orientation of the femoral neck associated with the acquisition of a bipedal gait in humans. Moreover, the precise determination of the three-dimensional orientation has implications in current research involved in developing clinical applications in diagnosis, hip surgery and rehabilitation.