Australopithecus afarensis : bipédie stricte ou associée à une composante arboricole ? Critiques et révision du matériel fémoral

Although much research has been carried out on Australopithecus afarensis locomotion, no consensus has yet been reached. Our new critic study on femoral material brings to the fore a strictly bipedal behaviour within this taxon. Our results are based on the pertinence of human anatomical characteristics among A. afarensis and on the absence of characteristics revealing arboreal displacement. These results have emerged from our different observation and interpretation of some preceding authors concerning the anatomy of these fossil hominids. It is important to underline that apomorphic characteristics of this species are difficult to interpret. They must not however be used to support the idea of arboreal displacement simply based on the fact of a no totally human morphe. We believe that present day humans do not necessarily reflect the earliest strict bipedal anatomic model. An the other hand, it appears that the disagreement between the two locomotor hypothesis for A. afarensis that are bipedalism and arboreal displacement, facing the possibility of bipedalism associated with negligible arboreal displacement, results more from an evolutionary fact than from a real scientific conflict.     

Neural control of quadrupedal and bipedal stance: implications for the evolution of erect posture

The transition among hominids from quadrupedalism to bipedalism resulted in modifications in their musculoskeletal morphology. It is unclear, however, whether changes in the circuitry of the CNS were also necessary in order to accommodate the unique balance requirements of two-limb support. This study addresses the issue of modifications in control strategies by investigating the rapid, automatic postural responses of feline and human subjects to sudden disturbances of balance in the anteroposterior (AP) direction while they stand quadrupedally and bipedally on movable platforms. Postural responses are characterized in terms of segmental adjustments, generated AP shear forces, and electromyographic activity. Feline and human subjects correct posture similarly when standing quadrupedally. Furthermore, both species correct stance primarily with their hindlimbs and use their forelimbs as supportive struts. In contrast, both species use completely different correctional strategies when standing bipedally. Morphological restrictions, however, prevent cats from adopting the pillar-like plantigrade posture of human beings. Thus, the correctional strategies of bipedal cats are distinct from those of bipedal human subjects. It is concluded that 1) automatic postural response patterns of quadrupedal Felis and bipedal Homo reflect the different biomechanical characteristics of the initial postures rather than species differences in CNS circuitry controlling stance; 2) hindlimb-dominated posture control is probably a common and relatively ancient pattern; and 3) reorganization of hominid CNS circuitry was probably unnecessary because hindlimb control was already a feature of the system.     

Palaeoenvironments and the origin of hominid bipedalism

Abstract

It has long been accepted that hominids emerged during the Pliocene in a savannah environment in which a terrestrial quadruped gradually developed bipedal adaptations. However, data from the Late Miocene (i.e. 7–7.5 Ma), including detailed palaeontological and biogeochemical studies, suggest that our earliest Upper Miocene ancestors inhabited well-wooded to forested environments where they could have spent a certain amount of time in the trees. A plausible type of ecosystem in which upright posture and bipedal locomotion could have emerged is represented by Miombo Woodland, in which vertical arboreal supports predominate and trees are separated from each other by gaps. Subsequently hominids dispersed into the Savannah as accomplished bipeds, but retained the ability to climb trees. This scenario is compatible with the postcranial anatomy of Australopithecus, including its femoral elongation, body proportions, manual precision grip (also present in 6-million-year-old Orrorin) and a non-prehensile hallux.     

The distal femoral anatomy of Australopithecus

The anatomy of the distal femoral fragments from Sterkfontein is reviewed, including its orthopaedic and biomechanical implications with respect to locomotion pattern. Comparisons are made with other hominids and a number of quadrupedal primates. Items which are considered are the obliquity and robustness of the shaft, the anterior intercondylar groove, the intercondylar notch, and the contour of the medial and lateral articular surfaces. The distinctive hominid status of these specimens is shown by their extensive adaptation to bipedal locomotion. No feature is found which is not fully commensurate with completely bipedal locomotion; rather, their distinctive hominid character points to a need for a reanalysis of the gait pattern in these early Pleistocene hominids.     

Palaeoenvironments and hominoid evolution

One of the key features that separates humans and their closest relatives (extinct species of the genus Homo and Praeanthropus and the australopithecines Australopithecus and Paranthropus) on the one hand, from the other hominoids, on the other, is their obligate bipedal locomotion when on the ground. This major difference from the generally quadrupedal locomotion practiced by other hominoids (Pan, Gorilla, Pongo and many extinct lineages) is reflected in many parts of the body, including all the major bones in the legs, arms, trunk and cranium. Locomotion has thus been of major interest to those interested in human origins, evolution, classification and phylogeny. A major hurdle to studies of the origins of bipedalism concerns the paucity of African hominoid fossils between 15 Ma, when all the adequately known hominoids were quadrupedal (most were pronograde, but at least one lineage was orthograde), and 4.2 Ma by which time fully bipedal hominids were established in Africa. Examination of Old World geology and palaeontology reveals a great deal about the evolution of palaeoenvironments and faunas during this period, and it is suggested that hominids evolved bipedal locomotion at the same time that there was a fundamental reorganisation of faunas towards the end of the Miocene. This faunal turnover resulted in the establishment of faunal lineages of "modern" aspect in Africa at the expense of "archaic" lineages which either went extinct or suffered a diminution of diversity. Many of the "modern" lineages were adapted to open country habitats in which grass became a major component of the diet as shown by modifications in the cheek teeth. Hominoids, in contrast, retained their traditional diet but were obliged to forage over greater and greater areas in order to do so, and this tactic led to pressures to modify the locomotor system rather than the diet. If bipedal hominids originated during this period, then the family Hominidae (sensu stricto) dates from about 8-7 Ma.     

La phalange distale du pouce d’Orrorin tugenensis (Miocène supérieur du Kenya)

The lion's share of articles dealing with the thumb anatomy of Plio-Pleistocene hominids has focussed on the capacity to manipulate and manufacture tools, and has largely neglected the locomotor aspects. However, in these hominids, the forelimb was still employed in locomotion. Certain of the anatomical characters classically associated with manipulation and/or fabrication of tools are already present in the Late Miocene species Orrorin tugenensis as shown by the terminal thumb phalanx BAR 1901'01. This specimen reveals crucial information suggesting that thumb morphology is not exclusively related to such tool using and manufacturing activities but reflects in a frequently bipedal creature a deeper adaptation concerning the precision grip essential for climbing and balancing, different from that of apes.     

Meat-eating by early hominids at the FLK 22Zinjanthropussite, Olduvai Gorge (Tanzania): an experimental approach using cut-mark data

The meat-eating behavior of Plio-Pleistocene hominids, responsible for the bone accumulations at the earliest archaeological sites, is still a hotly-debated issue in paleoanthropology. In particular, meat-eating and bone marrow consumption are often presented as either complementary or opposing strategies of carcass exploitation. The presence of cut marks on fossil archeofauna is a potential source of information that has not been consistently used as evidence of carcass consumption by hominids. Some authors interpret cut marks as the result of hominids manipulating meat-bearing bones, while others argue that they can also be the result of hominids extracting marginal scraps of carcass flesh that have survived carnivores’ initial consumption. In this study, a referential framework concerning the interpretation of cut marks is presented, based on a set of experiments conducted by the author. It is suggested, according to these experiments and data drawn from the FLK “Zinj” site, that hominids processed meat-bearing bones (on which flesh was abundant) rather than defleshed carcasses from felid kills.     

Physiology, thermoregulation and bipedalism

It has long been recognized that the bipedal posture reduces the surface area of the body exposed to the sun. In recent years, a theory has been developed by Wheeler that bipedalism evolved in the ancestor of the Hominidae in order to help relieve thermal stress on the animals in open equatorial environments. Bipedalism was said to afford a distinct adaptive advantage over quadrupedalism by permitting hominids to remain active in the open throughout the day. The heat load of the hypothetical hominid comprises the external environment as modelled by Wheeler and the animal's internal environment (i.e., the internal heat generated by its metabolic and locomotor activities, and its evaporative and respirative cooling capacities). When these factors are integrated in the calculation of the animal's thermal budget, the putative advantage of the bipedal over the quadrupedal posture is considerably reduced. The simulations conducted in this study suggest that the increased time afforded to early hominids in the open by bipedalism was relatively short and, therefore, of little or no adaptive significance. These results suggest that thermoregulatory considerations cannot be implicated as a first cause in the evolution of bipedalism in the hominid ancestor.     

New method of three-dimensional analysis of bipedal locomotion for the study of displacements of the body and body-parts centers of mass in man and non-human primates: Evolutionary framework

The current biomechanical interpretation of the chimpanzee's bipedal walking argues that larger lateral and vertical displacements of the body center of mass occur in the chimpanzee's “side-to-side” gait than in the human striding gait. The evolutionary hypothesis underlying this study is the following: during the evolution of human bipedalism one of the necessary changes could have been the progressive reduction of these displacements of the body center of mass. In order to quantitatively test this hypothesis, it is necessary to obtain simultaneously the trajectories of the centers of mass of the whole body and of the different body parts. To solve this problem, a new method of three-dimensional analysis of walking, associated with a volumetric modelling of the body, has been developed based on finite-element modelling. An orthogonal synchrophotographic device yielding four synchronous pictures of the walking subject allows a qualitative analysis of the photographic sequences together with the results of their quantitative analysis. This method was applied to an adult man, a 3-year-old girl and a 9-year-old male chimpanzee. Our results suggest that the trajectory of the body center of mass of the human is distinguished from that of the chimpanzee not by a lower movement amplitude but by the synchronization of the transverse and vertical displacements into two periodic curves in phase with one another. The non-human primate uses its repertoire of arboreal movements in its bipedal terrestrial gait, provisionally referred to as a “rope-walker” gait. We show that the interpretation of a “side-to-side” gait is not applicable to the chimpanzee. We argue that similarly this interpretation and the initial hypothesis presuppose a basic symmetric structure of the gait, in relation to the sagittal plane of progression, similar to the human one. This lateral symmetry of the right and left displacements of the center of gravity, in phase with the right and left single supports of walking, is probably a very derived feature of the human gait. We suggest that low lateral and vertical displacements of the body center of mass are not indicative of a progressive bipedal gait and we discuss the new evolutionary implications of our results. © 1993 Wiley-Liss, Inc.     

Arboreal bipedalism in wild chimpanzees: implications for the evolution of hominid posture and locomotion

Field observations of bipedal posture and locomotion in wild chimpanzees (Pan troglodytes) can serve as key evidence for reconstructing the likely origins of bipedalism in the last prehominid human ancestor. This paper reports on a sample of bipedal bouts, recorded ad libitum, in wild chimpanzees in Bwindi Impenetrable National Park in southwestern Uganda. The Ruhija community of chimpanzees in Bwindi displays a high rate of bipedal posture. In 246.7 hr of observation from 2001-2003, 179 instances of bipedal posture lasting 5 sec or longer were recorded, for a rate of 0.73 bouts per observation hour. Bipedalism was observed only on arboreal substrates, and was almost all postural, and not locomotor. Bipedalism was part of a complex series of positional behaviors related to feeding, which included two-legged standing, one-legged standing with arm support, and other intermediate postures. Ninety-six percent of bipedal bouts occurred in a foraging context, always as a chimpanzee reached to pluck fruit from tree limbs. Bipedalism was seen in both male and female adults, less frequently among juveniles, and rarely in infants. Both the frequency and duration of bipedal bouts showed a significant positive correlation with estimated substrate diameter. Neither fruit size nor nearest-neighbor association patterns were significantly correlated with the occurrence of bipedalism. Bipedalism is seen frequently in the Bwindi chimpanzee community, in part because of the unusual observer conditions at Bwindi. Most observations of bipedalism were made when the animals were in treetops and the observer at eye-level across narrow ravines. This suggests that wild chimpanzees may engage in bipedal behavior more often than is generally appreciated. Models of the likely evolutionary origins of bipedalism are considered in the light of Bwindi bipedalism data. Bipedalism among Bwindi chimpanzees suggests the origin of bipedal posture in hominids to be related to foraging advantages in fruit trees. It suggests important arboreal advantages in upright posture. The origin of postural bipedalism may have preceded and been causally disconnected from locomotor bipedalism.     

The biological and chronological maturation of early hominids

Recent studies on the rate and pattern of dental development indicate that the growth and maturation of early hominids were more similar to the extant apes than to modern humans. This contrasts with the previously held opinion derived from combined dental development, pattern and attrition studies claiming that early hominids were more hominine in their development (Mann, 1975). This paper explores the origin of this difference of opinion and reviews immature hominid dentitions with the benefit of improved radiographs and new data on the pattern and rate of pongid dental development. Paranthropus and Australopithecus specimens are shown to possess an ape-like development pattern but incisor development is specialized in the former and superficially human-like in pattern. The present and recent studies on dental development rate and pattern justify the position that early hominids were more ape-like in their growth and development. Therefore, ages at death calculated from pongid dental development schedules are provided for most immature early hominids. More detailed studies of early hominid developmental biology are now possible. It is suggested that divergent heterochronic processes characterize changes in brain/body proportions during hominid evolution. Relative rates of bone remodeling processes can now be identified on early hominid skeletons. The paleodemographic analysis of early hominids is little changed by the developmental model one chooses.     

Balance of the head in hominid evolution

In primates it is useful to distinguish three basic types of bipedal posture: (1) agonial, with extended hips and knees as in modern humans, (2) monogonial, with flexed hips but extended knees. and (3) digonial, with flexed hips and knees as in pongids. Early hominids retained an ancestral, forwardly inclined posture of the neck and head. Therefore the body posture of australopithednes must have differed from that in modem man, in which the centre of gravity of the head can be aligned with that of the body, other major centra of gravity, and important axes of rotation in a single frontal plane. It is suggested that in australopithednes the gravitational tilt of the head was counterbalanced by bent hips in association with hyperextended knees (monogonial posture). In australopithecines the increase in brain weight would have counteracted an improvement in the balance of the head. After the neck had assumed a more vertical posture as a consequence of shortening of the face, selection for an improved balance system in the bipedal posture favoured an increase in the weight of the postcondylar portion of the head, accentuated by selection for a posterior shift of the superior nuchal line in order to minimise the force of the nuchal muscles. At this stage the evolutionary increase in brain weight may have been largely a by-product of the process towards perfecting the bipedal posture. When the centre of gravity of the head had first become aligned with that of the body, the conditions of balance of the head had become favourable for a dramatic increase of brain size, as a result of selection for greater learning and storage capacity of the brain.     

制作工具在人类演化中的地位与作用

制作工具曾经被视作人类独有的行为能力,"人类"曾经据此而定义。但目前学术界将直立行走作为人类区别于其他灵长类最重要的体质与行为特征。少量其他动物种类,尤其是非人高等灵长类,也能使用工具乃至简单制作工具。如何认识制作工具在人类演化中的作用?人类制作工具的能力与其他动物有何区别?考古学是否有能力分辨人类的工具和其他灵长类的产品?本文通过对现代巴西猴群敲砸石头的行为及其产品、4300年前黑猩猩的"石制品"和早期人类石制品的比较研究,指出人类的工具与其他动物制作和使用的工具存在根本的区别;工具制作和使用对确定人类的演化方向,增强人类的适应生存能力,塑造人类的大脑与心智及行为方式,提升语言和交流能力,形成现代人类的身心和社会,至关重要,不可或缺。考古工作者一方面需要谨慎分辨、研究人类工具制作初期的产品,不使其与自然的产物和其他动作的作品相混淆,另一方面应该认识到人类工具制作在计划性、目的性、预见性、规范性和精美度上具有唯一性,有内在的智能控制、思维逻辑和规律可循。学科发展的积累和现代科技的支撑使考古学者具有多方面的利器,能够把人类工具制作的历史挖掘、复原出来,能够破译特定的石器技术和功能,进而将人类演化的历史画卷描绘得更加精细,更加完整。     

Pliocene hominid gait: New interpretations based on available footprint data from Laetoli

Normative trends in the gait patterns of modern man can be used to reconstruct crucial characteristics of the bipedal behaviour of Pliocene hominids from their fossilized footprints. In this reconstruction the interrelated parameters of velocity, stride-length, and cadence are determined from imprints made in damp volcanic ash some 3.7 million years ago. When early hominid footprint data is fitted to regression equations of high predictability for the interrelationship of these locomotor parameters in modern man, a pattern of gait emerges that contradicts previous reconstructions.     

Hominid paleoecology and competitive exclusion: Limits to similarity,niche differentiation,and the effects of cultural behavior

Fossil evidence from the Plio-Pleistocene of Africa apparently has confirmed a multi-lineage interpretation of early hominid evolution. Empirical refutation of the single species hypothesis must now be matched to the evolutionary ecology theory, which can underwrite taxonomic assessment and help to explain sympatric hominid coexistence. This paper contributes to that goal by reassessing the ecological rationale provided for the single-species hypothesis. Limiting similarity concepts indicate that the allowable ecological overlap between sympatric competitors is greater than the degrees of metric overlap often advanced as standards for identifying fossil species. Optimal foraging theory and the compression hypothesis show that the initial ecological reaction of a hominid to a sympatric competitor would likely be micro-habitat divergence and possibly also temporal differentiation of resource use. The long-term, evolutionary response is niche divergence, probably involving diet as well. General niche partitioning studies suggest that diet and habitat are the most common dimensions of niche separation, although temporal separation is unusually frequent in carnivores. The equation of niche with culture, basic to the single-species hypothesis, has no analytic meaning. Finally, four minor points are discussed, suggesting that (a) extinction is not unlikely, even for a long-lived and competitively competent hominid lineage, (b)parsimony is fickle, (c)interspecific mutualism may jeopardize survival, and (d)generalists are subordinate competitors, but for hominids, seemingly, successful ones. I argue that analog models of hominid paleoecology should be replaced by the use of zoological and anthropological observations to assess the generality and reliability of ecological theory and comcepts that may encompass early hominids.     

Senescence in European Middle Pleistocene hominids: The Atapuerca evidence

The mandibular specimens AT-250 and AT-793 (individual IV) and AT-888 (individual XXI) from the Atapuerca Middle Pleistocene sample show a combination of traits interpreted as evidence of senescence. In order to explore age-related mandibular changes relevant to the interpretation of the Atapuerca hominids, the sample of known age and sex from Spitalfields (London) and the Neolithic sample from Abu Hureyra (Syria) were used as a baseline. Results obtained indicate that the occurrence of an enlarged and posteriorly located mental foramen, in association with high alveolar resorption in AT-250 and AT-888, support the hypothesis of senescence in individuals IV and XXI from the Atapuerca sample. The findings suggest that European Middle Pleistocene populations lived long enough to reach senescence. The age at death for Middle Pleistocene hominids and Neanderthals has previously been estimated at around 40–45 years in the oldest specimens. The appearance of senescence processes before this age is indicative of higher rates of morphological aging in the clade which gave rise to Neanderthals. The anatomical significance of ageing features when considered in the context of craniofacial remodelling can help in the understanding of the taxonomic status of these morphological traits.     

Longitudinal study of dental development in chimpanzees of known chronological age: Implications for understanding the age at death of Plio-Pleistocene hominids

Reconstruction of life history variables of fossil hominids on the basis of dental development requires understanding of and comparison with the pattern and timing of dental development among both living humans and pongids. Whether dental development among living apes or humans provides a better model for comparison with that of Plio-Pleistocene hominids of the genus Australopithecus remains a contentious point. This paper presents new data on chimpanzees documenting developmental differences in the dentitions of modern humans and apes and discusses their significance in light of recent controversies over the human or pongid nature of australopithecine dental development. Longitudinal analysis of 299 lateral head radiographs from 33 lab-reared chimpanzees (Pan troglodytes) of known chronological age allows estimation of means and standard deviations for the age at first appearance of 8 developmental stages in the mandibular molar dentition. Results are compared with published studies of dental development among apes and with published standards for humans. Chimpanzees are distinctly different from humans in two important aspects of dental development. Relative to humans, chimpanzees show advanced molar development vis a vis anterior tooth development, and chimpanzees are characterized by temporal overlap in the calcification of adjacent molar crowns, while humans show moderate to long temporal gaps between the calcification of adjacent molar crowns. In combination with recent work on enamel incremental markers and CAT scans of developing dentitions of Plio-Pleistocene hominids, this evidence supports an interpretation of a rapid, essentially “apelike” ontogeny among australopithecines. © 1996 Wiley-Liss, Inc.     

Talocrural joint in African hominoids: implications for Australopithecus afarensis

Talocrural joints of the African apes, modern humans, and A.L.288-1 are compared in order to investigate ankle function in the Hadar hominids. Comparisons between the hominids and African pongids clearly illustrate the anatomical and mechanical changes that occurred in this joint as a consequence of the evolutionary transition to habitual bipedality. Features which are considered include the obliquity of the distal tibial articular surface, the shape of the talar trochlea, and the location and functional implications of the talocrural axis. In every functionally significant feature examined the A.L.288-1 talocrural joint is fully bipedal. Moreover, the Hadar ankle complex also shows the functional constraints which are necessarily imposed by the adaptation to habitual bipedalism.     

How did big brains evolve? The role of neonatal body size

The hypothesis that a mammalian species’ neonatal body size plays a role in determining its adult brain size is examined. This hypothesis is suggested by the observed correlation among mammals between neonatal body weight and adult brain weight. It is argued that there is no direct developmental linkage between these variables; rather, they are associated in evolution because of their opposing effects on the maturity level of the neonate. The evolution of neonate size in the hominids is also discussed, and consideration is given to trade-offs in pelvic design between locomotor and obstetrical functions. It is concluded that there was strong selection for brain enlargement in the hominids and that neonatal enlargement, rather than being intrinsically adaptive, was a direct response to the maturity-reducing effect of adult brain enlargement.