Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form. M ost research on the evolution of human locomotion has focused on walking. There are a few indications that the earliest-known hominids were bipeds 1,2 , and there is abundant fossil evidence that australopithecines habitually walked by at least 4.4 million years (Myr) ago However, although humans are comparatively poor sprinters, they also engage in a different type of running, endurance running (ER), defined as running many kilometres over extended time periods using aerobic metabolism. Although not extensively studied in non-humans, ER is unique to humans among primates, and uncommon among quadrupedal mammals other than social carnivores (such as dogs and hyenas) and migratory ungulates (such as wildebeest and horses) How well do humans run long distances? In considering human running, it helps to start from the perspective of the basic biomechanical differences that distinguish running and walking gaits in all mammals, including human bipeds. These differences are well characterized. Walking uses an 'inverted pendulum' in which the centre of mass vaults over a relatively extended leg during the stance phase, efficiently exchanging potential and kinetic energy out-of-phase with every ste
