UNIVERSITY OF CHICAGO PRESS, OCTOBER 2013But bipedality has evolved considerably since the first appearance of bipeds: it did not appear all at once. The awkward gait of the very primitive fossil hominin Ardipithecus ramidus (at 4.4 million years old, the earliest for which good skeletal evidence is known) shows that the first bipeds were not as refined as modern humans. They could stand upright, they could walk, though not as upright as modern humans, but they probably could not run very well. However, footprints attributed to the fossil human Australopithecus afarensis (Lucy) from a million years later show that by this time, creatures close to the human lineage could walk just about as well as modern humans. Even so, the skeleton of this creature was still very different from modern humans: Lucy could walk, but her skeleton suggests that she might have been a better tree climber than modern humans are.
The act of standing upright was followed, in sequence, by walking and then running—two gaits that demand very special, and rather different, adaptations. Daniel Lieberman and Dennis Bramble have recently proposed that many features of modern humans appear to be adaptations not to walking, as such, but to long-distance running. These include a range of features throughout the body not directly connected with the legs and feet.
Here are just two examples. Homo erectus and modern humans have barrel-shaped rib cages, in contrast to the cone-shaped, wide-bellied rib cages of earlier hominins. This means that later hominins had “waists,” which would have allowed the counterrotation of the arms relative to the legs while running. This is an extremely important aid to balance. Such counterrotation, however, ...