Locomotion in Alligator mississippiensis: kinematic effects of speed and posture and their relevance to the sprawling-to-erect paradigm

In terms of locomotory posture, amphibians and lizards are considered to be sprawlers, mammals and dinosaurs are considered to be erect, and extant crocodilians are considered to be intermediate because they use the 'high walk', a semi-erect posture where the body is held half-way between the sprawling and erect grades during locomotion. In addition, crocodilians occasionally use a sprawling posture. Extant crocodilians, therefore, provide an interesting model in which to investigate the sprawling-to-erect transition in vertebrate evolution. This study quantifies the sprawl and high walk kinematics of the alligator Alligator mississippiensis moving at different speeds on a treadmill and compares them with kinematic data available for other vertebrates. These data allow us to examine the effects of speed on crocodilian postures and to examine how crocodilian locomotion relates to the sprawling-to-erect paradigm in vertebrate locomotion. Our results show that the crocodilian sprawl is not functionally equivalent to the primitive sprawling behaviors exhibited by salamanders and lizards. In fact, although the high walks and sprawls of alligators exhibit some kinematic differences, they are actually much more similar than expected and, essentially, the crocodilian sprawl is a lower version of a high walk and could be termed a 'low walk'. In terms of the sprawling-to-erect transition, the high walk has knee kinematics intermediate between those of birds and non-archosaurian tetrapods, but alligators increase speed in a way completely different from other terrestrial vertebrates (distal rather than proximal limb elements are used to increase speed). These kinematic data viewed in the light of the fossil and phylogenetic evidence that modern crocodilians evolved from erect ancestors suggest that modern crocodilians have secondarily evolved a variable semi-erect posture and that they are problematic as an intermediate model for the evolutionary transition from sprawling to erect postures in archosaurs.