The evolution of locomotor stamina in tetrapods: circumventing a mechanical constraint

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Abstract
Endothermic tetrapods differ dramatically from ectothermic tetrapods in having a great capacity to sustain vigorous locomotion. I suggest that this difference reflects alternative adaptive responses to a mechanical constraint that was an inherent consequence of the vertebrate transition from aquatic to terrestrial modes of locomotion and respiration. The earliest tetrapods may not have been able to walk and breathe at the same time. Their sprawling gait and lateral vertebral bending would have required unilateral contractions of the thoracic musculature that may have interfered with the bilateral movements necessary for breathing. Modern lizards, whose locomotor and respiratory anatomy resembles that of the early tetrapods, provide support for this hypothesis because their breathing is greatly reduced during locomotor activity. Tetrapod lineages that gave rise to modern ectotherms apparently retained the constraint, becoming either highly specialized for burst activity based on anaerobic metabolism or specialized in passive mechanisms of defense against predators. The lineages from which birds and mammals are derived have undergone morphological changes that enable simultaneous running and breathing. In modern tetrapods upright posture is correlated with endothermic metabolism. This correlation may have arisen to circumvent ancestral constraints on locomotor stamina.