Locomotion in non-avian dinosaurs: integrating data from hindlimb kinematics, in vivo strains, and bone morphology

Analyses of non-avian dinosaur locomotion have been hampered by the lack of an appropriate locomotor analog among extant taxa. Birds, though members of the clade Dinosauria, have undergone significant modifications in hindlimb osteology and musculature. These changes have resulted in a uniquely developed system of limb kinematics (involving a more horizontal femoral posture and knee-dominated limb motion), which precludes the direct use of extant birds as models for non-avian dinosaur locomotion. Analyses of locomotor data from extant birds and mammals suggest a causal link between general hindlimb kinematics, bone strains, and limb bone morphology among these taxa. A model is proposed that relates the amount of torsional loading in femora to bone orientation, such that torsion is maximal in horizontal femora and minimal in vertical femora. Since bone safety factors are lower for torsional shear strains than for longitudinal axial strains, an increase in torsion can potentially affect bone morphology dramatically over evolutionary time. Interpreting the nearly identical limb bone dimensions and limb element proportions of non-avian dinosaurs and mammals in the light of this relationship supports the prediction of similar Vertical femoral postures and hip-driven limb kinematics in these two groups. This information can be used to interpret patterns of locomotor evolution within Dinosauria. The evolution of quadrupedalism with large body size and the acquisition of cursorial or graviportal limb morphologies occurred repeatedly but did not affect the underlying uniformity of dinosaur locomotor morphology. Only derived coelurosaurian theropods (paravians) developed significant modifications of the basic dinosaurian patterns of Limb use. Changes in theropod hindlimb kinematics and posture apparently began shortly prior to the origin of flight, but did not acquire a characteristically modern avian aspect until after the later acquisition of derived flight characteristics.