Caudofemoral musculature and the evolution of theropod locomotion

Living crocodilians and limbed lepidosaurs have a large caudofemoralis longus muscle passing from tail to femur. Anatomical and electromyographic data support the conclusion that the caudofemoralis is the principal femoral retractor and thus serves as the primary propulsive muscle of the hind limb. Osteological evidence of both origin and insertion indicates that a substantial caudofemoralis longus was present in archosaurs primitively and was retained in the clades Dinosauria and Theropoda. Derived theropods (e.g., ornithomimids, deinonychosaurs,Archaeopteryxand birds) exhibit features that indicate a reduction in caudofemoral musculature, including fewer caudal vertebrae, diminished caudal transverse processes, distal specialization of the tail, and loss of the fourth trochanter. This trend culminates in ornithurine birds, which have greatly reduced tails and either have a minute caudofemoralis longus or lack the muscle entirely.As derived theropod dinosaurs, birds represent the best living model for reconstructing extinct nonavian theropods. Bipedal, digitigrade locomotion on fully erect limbs is an avian feature inherited from theropod ancestors. However, the primitive saurian mechanisms of balancing the body (with a large tail) and retracting the limb (with the caudofemoralis longus) were abandoned in the course of avian evolution. This strongly suggests that details of the orientation (subhorizontal femur) and movement (primarily knee flexion) of the hind limb in extant birds are more properly viewed as derived, uniquely avian conditions, rather than as retentions of an ancestral dinosaurian pattern. Although many characters often associated with extant birds appeared much earlier in theropod evolution, reconstructing the locomotion of all theropods as completely birdlike ignores a wealth of differences that characterize birds.