Evolutionary reversals of limb proportions in early hominids? evidence from KNM-ER 3735 (Homo habilis)

Upper-to-lower limb proportions of Homo habilis are often said to be more ape-like than those of its reputed ancestor, Australopithecus afarensis. Such proportions would either imply multiple evolutionary reversals or parallel development of a relatively short upper limb in A. afarensis and later Homo. However, assessments of limb proportions are complicated by the fragmentary nature of the two known H. habilis skeletons, OH 62 and KNM-ER 3735. Initially, KNM-ER 3735 was compared to A.L. 288-1 (A. afarensis) using a single modern human and chimpanzee as reference. Here, based on a larger comparative sample, we find that the relative size of the distal humerus, radial head, and shaft of both KNM-ER 3735 and A.L. 288-1 lie within the range of variation of modern humans, whereas their sacra are small as is the case for all early hominids. In addition, their manual phalanges are similar in having a gracile base but robust midshaft. Contrary to earlier studies, the fossils are not differentiable from each other statistically with respect to all features listed above. On the other hand, they differ in robusticity of the scapular spine and relative length of the radial neck. An exact randomization test suggests only a very low probability of finding a similar degree of difference within a single species of extant hominoids. In contrast to the consensus view, we conclude that A.L. 288-1 had a short, human-like forearm, whereas KNM-ER 3735 possessed a distinctly longer forearm and more powerful shoulder girdle. This interpretation fits with earlier conclusions that suggested human-like humerofemoral proportions but chimpanzee-like brachial proportions for Homo habilis. Thus, the scenario of a unidirectional, progressive change in limb proportions within the hominid lineage is not supported by our work.