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Tortoises are famed for their slow locomotion, which is in part related to their herbivorous diet and the constraints imposed by their protective shells. For most animals, the metabolic cost of transport (CoT) is close to the value predicted for their body mass. Testudines appear to be an exception to this rule, as previous studies indicate that, for their body mass, they are economical walkers. The metabolic efficiency of their terrestrial locomotion is explainable by their walking gait biomechanics and the specialisation of their limb muscle physiology, which embodies a predominance of energy-efficient slow-twitch type I muscle fibres. However, there are only two published experimental reports of the energetics of locomotion in tortoises, and these data show high variability. Here, Mediterranean spur-thighed tortoises (Testudo graeca) were trained to walk on a treadmill. Open-flow respirometry and high-speed filming were simultaneously used to measure the metabolic cost of transport and to quantify limb kinematics, respectively. Our data support the low cost of transport previously reported and demonstrate a novel curvilinear relationship to speed in Testudines, suggesting tortoises have an energetically optimal speed range over which they can move in order to minimise the metabolic cost of transport. 

Abstract Armoured, rigid bodied animals, such as Testudines, must self-right should they find themselves in an inverted position. The ability to self-right is an essential biomechanical and physiological process that influences survival and ultimately fitness. Traits that enhance righting ability may consequently offer an evolutionary advantage. However, the energetic requirements of self-righting are unknown. Using respirometry and kinematic video analysis, we examined the metabolic cost of self-righting in the terrestrial Mediterranean spur-thighed tortoise and compared this to the metabolic cost of locomotion at a moderate, easily sustainable speed. We found that self-righting is, relatively, metabolically expensive and costs around two times the mass-specific power required to walk. Rapid movements of the limbs and head facilitate successful righting however, combined with the constraints of breathing whilst upside down, contribute a significant metabolic cost. Consequently, in the wild, these animals should favour environments or behaviours where the risk of becoming inverted is reduced.