Innovations permit the diversification of lineages, but they may also impose functional constraints on behaviors such as locomotion. Thus, it is not surprising that secondary simplification of novel locomotory traits has occurred several times among vertebrates and could potentially lead to exceptional divergence when constraints are relaxed. For example, the gecko adhesive system is a remarkable innovation that permits locomotion on surfaces unavailable to other animals, but has been lost or simplified in species that have reverted to a terrestrial lifestyle. We examined the functional and morphological consequences of this adaptive simplification in the Pachydactylus radiation of geckos, which exhibits multiple unambiguous losses or bouts of simplification of the adhesive system. We found that the rates of morphological and 3D locomotor kinematic evolution are elevated in those species that have simplified or lost adhesive capabilities. This finding suggests that the constraints associated with adhesion have been circumvented, permitting these species to either run faster or burrow. The association between a terrestrial lifestyle and the loss/reduction of adhesion suggests a direct link between morphology, biomechanics, and ecology.biomechanics | toepads | Pachydactylus | adaptation | Namibia S tructural innovations involving coordinated changes in multiple anatomical systems [Frazzetta's "complex adaptations" (1)] are associated with the diversification of many groups of vertebrates. Many such innovations often occur repeatedly within, as well as between, clades. One example is the evolution of the prehensile tail, which has arisen in primates, nonprimate mammals, seahorses, amphibians, and several groups of lizard, allowing its possessors to move through the environment in novel ways (2-6). Although the acquisition of such innovations is often implicated in both diversification and ecological specialization, much less is known about the causes and consequences of their secondary reduction and loss. If such novelties promote organismal diversity, then the evolutionary simplification of such innovations might commonly occur only when their diminution becomes even more advantageous. Generally, across the tree of life, we might expect that many of the most extraordinary examples of adaptive divergence should coincide not only with the origin and retention of innovations but also with their simplification, and ultimately their loss, in association with adaptation to new situations.Much of what is known about the secondary simplification of locomotor structures relates to limb loss/reduction in tetrapods (7-11), such as in snakes and lizards. The increase in the rate of evolutionary change in these situations suggests that the origin of an elongate body likely led not only to relaxed selection for the retention of limbs, but also to rapid adaptive selection, favoring both their loss and other associated morphological changes. In these cases, it is expected that both the rate of morphological evolution and the area...