Was “Lucy” more human than her “child”? Observations on early hominid postcranial skeletons

Fully adult partial skeletons attributed to Australopithecus afarensis (AL 288-1, “Lucy”) and to Homo habilis (OH 62, “Lucy's child”), respectively, both include remains from upper and lower limbs. Relationships between various limb bone dimensions of these skeletons are compared to those of modern African apes and humans. Surprisingly, it emerges that OH 62 displays closer similarities to African apes than does AL 288-1. Yet A. afarensis, whose skeleton is dated more than 1 million years earlier, is commonly supposed to be the ancestor of Homo habilis. If OH 62, classified as Homo habilis by its discoverers, does indeed represent a stage intermediate between A. afarensis and later Homo, a revised interpretation of the course of human evolution would be necessary.     

Size- and locomotion-related aspects of hominid and anthropoid pelves: An osteometrical multivariate analysis

Two multivariate methods — the logarithmic principal component analysis (LPCA), and the logarithmic factorial analysis (LFA) — have been used tocompare the hip bone proportions of hominoids biometrically. The results have shown that size effects among apes and hominids interact to a centain extent with locomotor specializations, which are related to the attainment of more or less terrestrial behaviors. The pelvic morphology of great apes (Pongo, Pan, Gorilla) has retained numerous morphological traits — such as a gracile and elongated hip bone —, which were inherited from common adaptations to arboreal locomotion. In spite of these common traits, the African pongids (Pan, Gorilla) present two very different pelvic morphologies corresponding to two adaptative modes of terrestrial quadrupedalism. The hip bone of humans is proportionnally short and robust, most particularly at the level of its axial part. These characteristics, as well as the whole pelvic proportions, clearly indicate that gravitational forces exert a strong pressure on the pelvic walls during bipedalism. Among hominids, the transition from an australopithecine-like pelvic pattern to a human-like one corresponds to an increase of loading constraints on the hip jiont. This seems to indicate an evident change in locomotor behavior. Progression apparently became exclusively terrestrial with the genusHomo.     

Segment and joint angles of hind limb during bipedal and quadrupedal walking of the bonobo (Pan paniscus)

We describe segment angles (trunk, thigh, shank, and foot) and joint angles (hip, knee, and ankle) for the hind limbs of bonobos walking bipedally ("bent-hip bent-knee walking," 17 sequences) and quadrupedally (33 sequences). Data were based on video recordings (50 Hz) of nine subjects in a lateral view, walking at voluntary speed. The major differences between bipedal and quadrupedal walking are found in the trunk, thigh, and hip angles. During bipedal walking, the trunk is approximately 33-41 degrees more erect than during quadrupedal locomotion, although it is considerably more bent forward than in normal human locomotion. Moreover, during bipedal walking, the hip has a smaller range of motion (by 12 degrees ) and is more extended (by 20-35 degrees ) than during quadrupedal walking. In general, angle profiles in bonobos are much more variable than in humans. Intralimb phase relationships of subsequent joint angles show that hip-knee coordination is similar for bipedal and quadrupedal walking, and resembles the human pattern. The coordination between knee and ankle differs much more from the human pattern. Based on joint angles observed throughout stance phase and on the estimation of functional leg length, an efficient inverted pendulum mechanism is not expected in bonobos.     

Energetic costs of bipedal and quadrupedal walking in Japanese macaques

We investigated the energetic costs of quadrupedal and bipedal walking in two Japanese macaques. The subjects were engaged in traditional bipedal performance for years, and are extremely adept bipeds. The experiment was conducted in an airtight chamber with a gas analyzer. The subjects walked quadrupedally and bipedally at fixed velocities (<5 km/hr) on a treadmill in the chamber for 2.5-6 min. We estimated energy consumption from carbon dioxide (CO2) production. While walking bipedally, energetic expenditure increased by 30% relative to quadrupedalism in one subject, and by 20% in another younger subject. Energetic costs increased linearly with velocity in quadrupedalism and bipedalism, with bipedal/quadrupedal ratios remaining almost constant. Our experiments were relatively short in duration, and thus the observed locomotor costs may include presteady-state high values. However, there was no difference in experimental duration between bipedal and quadrupedal trials. Thus, the issue of steady state cannot cancel the difference in energetic costs. Furthermore, we observed that switching of locomotor mode (quadrupedalism to bipedalism) during a session resulted in a significant increase of CO2 production. Taylor and Rowntree ([1973] Science 179:186-187) noted that the energetic costs for bipedal and quadrupedal walking were the same in chimpanzees and capuchin monkeys. Although the reason for this inconsistency is not clear, species-specific differences should be considered regarding bipedal locomotor energetics among nonhuman primates. Extra costs for bipedalism may not be great in these macaques. Indeed, it is known that suspensory locomotion in Ateles consumes 1.3-1.4 times as much energy relative to quadrupedal progression. This excess ratio surpasses the bipedal/quadrupedal energetic ratios in these macaques.     

Defence of bipedalism

The long-unresolved and much publicized puzzle of how human ancestors could have ventured upon habitual bipedalism without making themselves highly vulnerable to predation derives from a misunderstanding of the principles upon which predator-prey relations work. Erect stature, regardless of locomotion proficiency, in fact enhances passive, and not simply active, defence against predators. This was the necessary condition, the security ‘cover’, for selection for the elaboration of bipedal behavior. As illustration, I analyse the defence systems ofPan troglodytes and the hunting techniques of their predators, known and potential, in order to hypothesize the defence impact of bipedal selection on the former; the relevance of human experience with felid predators, in particular, is also then explored. Despite known major differences in the late Miocene mammalian predator guild, the predator-prey framework is likely to have been much the same as to-day, making the findings applicable to the situation of proto-hominids. Once it is accepted selection pressure for bipedalism was originally focused on posture and not locomotion, the mystery of its emergence disappears.     

Laetoli toes andAustralopithecus afarensis

The probable misfit between feet, particularly toes II–V, of 3.0-million-year-oldAustralopithecus afarensis from Hadar, Ethiopia, and the 3.5-million-year-old hominid footprints at Site G, Laetoli, Tanzania, casts doubt thatA. Afarensis made the Laetoli trails. We suggest that another species ofAustralopithecus or an anonymous genus of the Hominidae, with remarkably humanoid feet, walked at Laetoli. It would be imprudent to declare thatHomo was present at Laetoli 3.5 million years ago (my) because there is no evidence of brain expansion, advanced tool manufacture, or other non-locomotor hallmarks of the human condition at Site G.     

Three-dimensional kinematics of capuchin monkey bipedalism

Capuchin monkeys are known to use bipedalism when transporting food items and tools. The bipedal gait of two capuchin monkeys in the laboratory was studied with three-dimensional kinematics. Capuchins progress bipedally with a bent-hip, bent-knee gait. The knee collapses into flexion during stance and the hip drops in height. The knee is also highly flexed during swing to allow the foot which is plantarflexed to clear the ground. The forefoot makes first contact at touchdown. Stride frequency is high, and stride length and limb excursion low. Hind limb retraction is limited, presumably to reduce the pitch moment of the forward-leaning trunk. Unlike human bipedalism, the bipedal gait of capuchins is not a vaulting gait, and energy recovery from pendulum-like exchanges is unlikely. It extends into speeds at which humans and other animals run, but without a human-like gait transition. In this respect it resembles avian bipedal gaits. It remains to be tested whether energy is recovered through cyclic elastic storage and release as in bipedal birds at higher speeds. Capuchin bipedalism has many features in common with the facultative bipedalism of other primates which is further evidence for restrictions on a fully upright striding gait in primates that transition to bipedalism. It differs from the facultative bipedalism of other primates in the lack of an extended double-support phase and short aerial phases at higher speeds that make it a run by kinematic definition. This demonstrates that facultative bipedalism of quadrupedal primates need not necessarily be a walking gait.