The Anatomical and Functional Evolution of the Femoral Fourth Trochanter in Ornithischian Dinosaurs

Abstract

The femoral fourth trochanter is the attachment site of the caudofemoralis musculature, which is the primary hindlimb retractor in most non-avian dinosaurs. Early ornithischian dinosaurs are uniquely characterized by a fourth trochanter with a prominent pendant process. Throughout the evolution of ornithischians, the fourth trochanter repeatedly converged on two major morphological changes: (1) the distal migration of the trochanter down the femoral shaft and (2) the loss of the pendant process. Both changes, as well as the original evolution of the pendant form, relate to a single major functional shift emphasizing caudofemoral leverage. Direct evidence of muscle scarring across the surface of the pendant process affirms that it served to extend the attachment of the primary caudofemoralis brevis tendon distally. A proximally located fourth trochanter is the basal condition in dinosaurs and other archosaurs, and the development of a pendant process lengthened the functional lever arm with regard to the insertion of the caudofemoralis. This adaptation afforded improved mechanical advantage, perhaps beneficial in the context of the newly assumed herbivorous diets of basal ornithischians. As some derived ornithischians increased in body size, a high-leverage system with a more distal caudofemoralis attachment evolved. In some groups, the fourth trochanter as a whole descended down the femur, eventually reaching a point where the pendant process was unnecessary. Sauropodomorphs, the other great lineage of dinosaur herbivores, converged on the same high-leverage distal fourth trochanter arrangement, but without first transitioning through a prominent pendant form. Anat Rec, 303:1146-1157, 2020. © 2019 Wiley Periodicals, Inc.