March of the Titans: The Locomotor Capabilities of Sauropod Dinosaurs

Sauropod dinosaurs are the largest terrestrial vertebrate to have lived on Earth. This size must have posed special challenges for the musculoskeletal system. Scaling theory shows that body mass and hence the loads that must be overcome increases with body size more rapidly than either the ability of the muscles to generate force, or the ability of the skeleton to support these loads. Here we demonstrate how one of the very largest sauropods, Argentinosaurus huinculensis (40 metres long, weighing 83 tonnes), may have moved. A musculoskeletal model was generated using data captured by laser scanning a mounted skeleton and assigning muscle properties based on comparative data from living animals. Locomotion is generated using forward dynamic simulation to calculate the accelerations produced by the muscle forces, coupled with machine learning techniques to find a control pattern that minimises metabolic cost. The simulation demonstrates that at such vast body size, joint range of motion needs to be restricted to allow sufficient force generation for an achievable muscle mass. However when this is done, a perfectly plausible gait can be generated relatively easily. Whilst this model represents the best current simulation of the gait of these giant animals, it is likely that there are as yet unknown mechanical mechanisms, possibly based on passive elastic structures that should be incorporated to increase the efficiency of the animal's locomotion. It is certainly the case that these would need to be incorporated into the model to properly assess the full locomotor capabilities of the animal.

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