Acquisition of bipedalism: the Miocene hominoid record and modern analogues for bipedal protohominids.

The well-known fossil hominoid Proconsul from the Early Miocene of Kenya was a non-specialized arboreal quadruped with strong pollicial/hallucial assisted grasping capability. It lacked most of the suspensory specializations acquired in living hominoids. Nacholapithecus, however, from the Middle Miocene of Kenya, although in part sharing with Proconsul the common primitive anatomical body design, was more specialized for orthograde climbing, 'hoisting' and bridging, with the glenoid fossae of the scapula probably being cranially orientated, the forelimbs proportionally large, and very long toes. Its tail loss suggests relatively slow movement, although tail loss may already have occurred in Proconsul. Nacholapithecus-like positional behaviour might thus have been a basis for development of more suspensory specialized positional behaviour in later hominoids. Unfortunately, after 13 Ma, there is a gap in the hominoid postcranial record in Africa until 6 Ma. Due to this gap, a scenario for later locomotor evolution prior to the divergence of Homo and Pan cannot be determined with certainty. The time gap also causes difficulties when we seek to determine polarities of morphological traits in very early hominids. Interpretation of the form-function relationships of postcranial features in incipient hominids will be difficult because it is predicted that they had incorporated bipedalism only moderately into their total positional repertoires. However, Japanese macaques, which are trained in traditional bipedal performance, may provide useful hints about bipedal adaptation in the protohominids. Kinematic analyses revealed that these macaques walked bipedally with a longer stride and lower stride frequency than used by ordinary macaques, owing to a more extended posture of the hindlimb joints. The body centre of gravity rises during the single-support phase of stance. Energetic studies of locomotion in these bipedal macaques revealed that energetic expenditure was 20-30% higher in bipedalism than in quadrupedalism, regardless of walking velocity.