The femur of Orrorin tugenensis exhibits morphometric affinities with both Miocene apes and later hominins

Orrorin tugenensis (Kenya, ca. 6 Ma) is one of the earliest putative hominins. Its proximal femur, BAR 1002000, was originally described as being very human-like, although later multivariate analyses showed an australopith pattern. However, some of its traits (for example, laterally protruding greater trochanter, medially oriented lesser trochanter and presence of third trochanter) are also present in earlier Miocene apes. Here, we use geometric morphometrics to reassess the morphological affinities of BAR 1002'00 within a large sample of anthropoids (including fossil apes and hominins) and reconstruct hominoid proximal femur evolution using squared-change parsimony. Our results indicate that both hominin and modern great ape femora evolved in different directions from a primitive morphology represented by some fossil apes. Orrorin appears intermediate between Miocene apes and australopiths in shape space. This evidence is consistent with femoral shape similarities in extant great apes being derived and homoplastic and has profound implications for understanding the origins of human bipedalism.