Burly gaits: centers of mass, stability, and the trackways of sauropod dinosaurs

The narrow- and wide-gauge trackways attributed to sauropod dinosaurs are hypothesized to be a consequence of the relative positions of their centers of mass. This hypothesis was tested using three-dimensional, trackway-producing computer models of two sauropods and studies of Asian elephants. Centers of mass of sauropod models were computed using density distributions that reflect the high degree of pneumatization of the skeletons and air sacs within the body. A close correspondence was found between the relative areas of hand and foot prints in different trackways and the relative fractions of the body weight borne by the forefeet and hindfeet in the different types of sauropods inferred to have made the trackways. Experimental studies of Asian elephants corroborated the close correspondence between relative areas of the hindfeet and forefeet and body weight distribution. Replicating actual sauropod trackways with the walking models enabled testing of proposed gaits for a sauropod model. Brachiosaurus brancai, with its more centrally positioned center of mass, was stable and possessed a wide safety margin only when replicating a wide trackway. Conversely, Diplodocus carnegii, with a more posteriorly placed center of mass, was most stable when replicating a narrow trackway. A trend for large sauropods (>12 tons), independent of clade, to have more anteriorly positioned centers of mass was identified, and it is proposed that all large sauropods were restricted to producing wide-gauge trackways for stability reasons. The primitive gait state for Sauropodomorpha was determined to be one that produced narrow-gauge tracways.