In this paper I look at the scaling of the energy costs of locomotion, and ask whether we can explain what we observe. The explanations must depend on mathematical models. If we cannot formulate a convincing model that predicts a scaling rule reasonably accurately, we have failed to explain the rule. The reverse is unfortunately not true; a model that predicts a scaling rule correctly does not guarantee that our explanation is correct, because several models may predict the same rule.My interest here is in widely applicable scaling rules; for example in rules that will predict the scaling of running over the range from small rodents to elephants, or of flight from sparrows to swans. Mice are not scale models of elephants, and do not move like tiny elephants, and sparrows are not miniature swans. The models will have to be very general, incorporating little specific anatomical or kinematic detail. Conveniently, this implies that they will be simple. Many simple models of running, swimming and flight were presented at the first Scaling Conference (Pedley, 1977), on which this paper builds.In the decades preceding the first Scaling Conference, measurements of metabolic rate during locomotion had been greatly facilitated by the introduction of methods using treadmills (Taylor et al., 1970), wind tunnels (Tucker, 1968) and water tunnels (Brett, 1964). Allometric exponents relating the measured energy cost of locomotion to body mass had been calculated by Taylor et al. (1970) for running; by Tucker (1970) for running and flight; and for swimming by SchmidtNielsen (1972). Allometric equations in more recent papers are referred to in later sections of this one.Because muscles do not work with uniform efficiency, it is much more difficult to devise a model that predicts the metabolic energy cost of locomotion than one that predicts mechanical work. In contrast, oxygen consumption (and hence metabolic power) can be measured directly, whereas determination of mechanical work in locomotion generally involves calculations subject to a good deal of uncertainty. Thus comparisons between theoretical and observed energy costs are not easy. It is the metabolic cost of locomotion, rather than the mechanical work, which is important for the animal's energy budget.Dynamic similarity I will refer frequently to the concept of dynamic similarity To achieve the required generality, models designed to predict scaling relationships for diverse groups of animals generally need to be simple. An argument based on considerations of dynamic similarity predicts correctly that the mechanical cost of transport for running [power/(body mass ϫ speed)] will be independent of body mass; but measurements of oxygen consumption for running birds and mammals show that the metabolic cost of transport is proportional to (body mass) −0.32 . Thus the leg muscles seem to work more efficiently in larger animals. A model that treats birds as fixed wing aircraft predicts that the mechanical power required for flight at the maximum range speed will be proportiona...