Bipedal behavior of olive baboons (Papio anubis) and its relevance to an understanding of the evolution of human bipedalism

The bipedal behavior of a troop of olive baboons (Papio anubis) is described. Bipedalism is relatively rare but nevertheless occurs in a wide variety of situations, although bipedalism during feeding occurs much more frequently than in other situations. The incidence of bipedalism varies between different age-sex classes and between individuals within age-sex classes. This pattern of bipedalism occurred within an overall adaptive response, particularly in feeding behavior, which was similar to that of the gelada baboon (Theropithecus gelada). The data on bipedalism is used together with an existing model of early hominid differentiation based on T. gelada to indicate the types of bipedal behavior which might have occurred in early hominid small object feeders and to suggest how a bipedal pattern of this type might have served as a basis for the action of selection for a more committedly bipedal pattern at later stages of hominid evolution.     

A new pelvic fragment from swartkrans and the relationship between the robust and gracile australopithecines

A recently discovered hominid pelvic fragment from Swartkrans (SK 3155) is described in detail with particular reference to the relationship of the two presently recognized forms of australopithecines in South Africa. Results of this examination and metrical analysis indicate that the acetabulum and iliac blade of the early hominids are similar to Homo sapiens except for a unique pattern of traits: a relatively small sacral articular surface, a relatively small acetabulum, a relatively large iliac fossa, and wide lateral splaying of the iliac blades. The new Swartkrans fossil expresses these traits more strongly than does the gracile australopithecine (Sts 14) and is therefore somewhat less similar to Homo sapiens but it is very unlike any pongid.     

The relative cost of bent-hip bent-knee walking is reduced in water

The debate about how early hominids walked may be characterised as two competing hypotheses: They moved with a fully upright (FU) gait, like modern humans, or with a bent-hip, bent-knee (BK) gait, like apes. Both have assumed that this bipedalism was almost exclusively on land, in trees or a combination of the two. Recent findings favoured the FU hypothesis by showing that the BK gait is 50–60% more energetically costly than a FU human gait on land. We confirm these findings but show that in water this cost differential is markedly reduced, especially in deeper water, at slower speeds and with greater knee flexion. These data suggest that the controversy about australopithecine locomotion may be eased if it is assumed that wading was a component of their locomotor repertoire and supports the idea that shallow water might have been an environment favourable to the evolution of early forms of “non-optimal” hominid bipedalism.     

Heterochronic processes in human evolution: An ontogenetic analysis of the hominid pelvis

Changes in pelvic shape in human ontogeny and hominid phylogeny suggest that the heterochronic processes involved differ greatly from the neotenic process traditionally described in the evolution of the skull. The morphology of 150 juvenile and adult pelves of African apes, 60 juvenile and adult pelves of modern humans, two adult pelves and a juvenile hip bone of australopithecines (Sts 14, AL 288, MLD 7) was studied. Multivariate results, ontogenetic allometries, and growth curves confirm that the pelvic growth pattern in humans differs markedly from those of the African apes. The results permit the following conclusions. First, the appearance of a new feature (acetabulo-cristal buttress and cristal tubercle) at the time of human birth allows the addition of traits, such as the attainment of a proportionally narrower pelvis, with more sagittally positioned iliac blades. Pelvic proportions and orientation change progressively in early childhood as bipedalism is practiced. Other changes in pelvic proportions occur later with the adolescent growth spurt. Second, comparison of juvenile and adult australopithecines to modern humans indicates that 1) some pelvic traits of adult Australopithecus resemble those of neonate Homo; 2) the pelvic growth of Australopithecus was probably closer to that of apes, than to that of humans; and 3) prolonged growth in length of hindlimb and pelvis after sexual maturity seems to be a unique feature of Homo. The position of the acetabulo-cristal buttress and of the cristal tubercle on the ilium are similar in adult Australopithecus and neonate Homo suggesting that this feature may have been displaced later during hominid evolution. Progressive displacement of the acetabulo-cristal buttress on the ilium occurs both during hominid evolution (from Australopithecus to Homo sapiens) and human growth (from neonate to adult). This suggests peramorphic evolution of the pelvic morphology of hominids combining three processes of recapitulation (pre-displacement, acceleration and time hypermorphosis). The results lend credence to the hypothesis that no single heterochronic process accounts for all human evolutionary change; rather this reflects a combination of relative changes in growth rhythm and duration, including other perturbations, such as the appearance of new morphological features. Am J Phys Anthropol 105:441–459, 1998. © 1998 Wiley-Liss, Inc.     

The ischium and hip extensor mechanism in human evolution

Although it is commonly stated that the ischia of the late Pliocene–early Pleistocene hominid fossils are long and ape-like, new interpretations show this view to be fallacious. An important new theory proposed by Robinson concludes that the gracile form of early hominid was an efficient biped, but the robust form was a less efficient biped and was adapted for tree climbing. Interpretation of the ischium is crucial to this idea. The present study shows that (1) the gracile and robust australopithecine ischia had similar relative lengths and (2) that the hamstring mechanism was probably very similar in the two forms of South African early hominid.     

A new morphometric analysis of the hominid pelvic bone

This study is based upon a new morphometric technique providing both size and shape variables. It has been applied to 189 pelvic bones of extant humans and African apes as well as to 13 hominid pelvic bones of various taxonomic status. The main aim of this work is to include such fossil bones in the same study in order to set a synthetic comparison of their shape in the light of the yardstick given by the African ape/human pelvic bone comparison. To do so, ratio diagrams are chosen because they are simple and very expressive tools with which to present such comparisons. Shape differences are very well illustrated and quantified by this technique. The ilium appears to be the most different of the three parts of the pelvic bone. Compared to these differences, discrepancies between fossil hominid and extant human bones are of a totally different scale. This shows the architectural unity related to the acquisition of bipedalism by hominids. It is nonetheless possible to detect two levels of difference. The first separates Australopithecus from Homo and could be seen as reflecting locomotor differences between both genera. The second splits both Homo erectus and Neanderthal from modern human pelvic bones. It appears from the hominid fossil record of pelvic bones that two periods of stasis exist and are separated by a period of very rapid evolution corresponding to the emergence of the genus Homo. We are of the opinion that the same could be true for the split between African ape and hominid lineages at the end of the Miocene.     

Affinities of the Swartkrans 847 hominid cranium

Further evidence of the presence of a second hominid species at the Swartkrans locality was obtained in 1969 when the SK.847 specimen was discovered by us to represent the same individual as the SK.80 maxilla. The SK.847 specimen had previously been regarded as robust australopithecine, whereas the latter was first attributed to Telanthropus capensis and subsequently to a species of the genus Homo. Recent criticism of our interpretation of these remains has not evaluated and analyzed critically the primary fossil evidence. Instead it relies on a strict adherence to an as yet unsubstantiated hypothesis that posits only a single hominid species at any point in space and time in the Cenozoic history of Hominidae.     

Carrying, Sharing, and Hand Preference in Tufted Capuchins (Cebus apella)

We examined the relationship among carrying, food-sharing, and hand preference in tufted capuchins (Cebus apella). The rationale was to evaluate further the use of Cebus as an alternative primate model to Pan for behavior relevant to early hominid evolution. We first examined bipedalism and food-sharing within an established social group, and then examined the direction and strength of hand preference for food carrying in an expanded sample. Several aspects of capuchin behavior warrant discussion. First, bipedal carrying and food-sharing occurred more frequently when we provided bulky foods than when we provided smaller foods. Second, food-sharing was characterized by passive tolerance, rather than active giving, between subjects. Third, subjects shared food primarily with immatures and followed a pattern of reciprocal exchange. Finally, we found no evidence for population-level hand preference for carrying. We posit that an array of behavioral similarities among Cebus, Pan, and Homo evolved through convergent processes, and in this regard capuchins can be seen as an alternative primate model to chimpanzees for the evolution of early hominid behavior.     

Biomechanical interpretation of the early hominid hip

A new pelvic fragment from Swartkrans provides the opportunity to analyze the hip joint mechanics of the robust form of early hominid. The function of the lateral support system provided by the abductor muscles of the hip appears to be similar to that of the gracile early hominid from Sterkfontein. The system is well adapted for providing the lateral support necessary for efficient bipedalism. The hip extensor mechanism and hip internal rotatory system also appear to be well adapted for efficient bipedalism in a way very similar to the other early hominids. The conclusion reached is that the robust and gracile forms of South African early hominids were basically similar in their locomotor adaptation and were most likely habitual bipeds.     

African ape ancestry

It is commonly believed that the australopithecines are more closely related to humans than to African apes. This view is hardly compatible with the biomolecular data which place theHomo/Pan split at the beginning of the australopithecine period. Nothing in the fossil hominid morphology precludes the possibility that some australopithecines were ancestral to gorillas or chimpanzees and others to humans.     

History of American paleoanthropological research on early Hominidae, 1925–1980

Understanding of the early stages of hominid evolution prior to 1925 was based primarily on comparative morphological evidence derived from extant primates. With the publication of Australopithecus by Dart in 1925 and subsequent research in South Africa, new possibilities for empirical assessment of early hominid evolutionary history were opened. It was Gregory's work, with Hellman, reported at the first meeting of the AAPA in 1930, that convinced many workers of the hominid status of Australopithecus. The debunking of Eoanthropus as a Pliocene hominid, far from having a totally negative effect, showed that cranial expansion had occurred after bipedalism in hominid evolution, demonstrated that chemical dating had come of age, and in a broader sense, had underlined that phylogenetic hypotheses are falsifiable by recourse to the evidence. The input of biological sciences into early hominid studies, as exemplified by Washburn's “new physical anthropology,” reduced taxonomic diversity and focused attention on paleoecology and behavior. The development of the multidisciplinary approach to field research, pioneered by L. Leakey and brought to fruition by Howell, was of fundamental importance in accurately dating and understanding the context of early hominids. Archaeology, primatology, comparative and functional morphology, and morphometrics have contributed substantially in recent years to a fuller understanding of early hominid evolution. American granting agencies have heavily supported early hominid research but patterns of funding have not kept pace with the change from research based largely on individualistic enterprise to multidisciplinary research projects. Future early hominid research, if funding is available, will likely be directed toward investigating temporal and geographic gaps now known in the fossil record and in more rigorous and multidisciplinary investigations of early hominid behavior.     

Dental metric assessment of the omo fossils: implications for the phylogenetic position of Australopithecus africanus

The discovery of Australopithecus afarensis has led to new interpretations of hominid phylogeny, some of which reject A. africanus as an ancestor of Homo. Analysis of buccolingual tooth crown dimensions in australopithecines and Homo species by Johanson and White (Science 202:321-330, 1979) revealed that the South African gracile australopithecines are intermediate in size between Laetoli/hadar hominids and South African robust hominids. Homo, on the other hand, displays dimensions similar to those of A. afarensis and smaller than those of other australopithecines. These authors conclude, therefore, that A. africanus is derived in the direction of A. robustus and is not an ancestor of the Homo clade. However, there is a considerable time gap (ca. 800,000 years) between the Laetoli/Hadar specimens and the earliest Homo specimens; "gracile" hominids from Omo fit into this chronological gap and are from the same geographic area. Because the early specimens at Omo have been designated A. afarensis and the later specimens classified as Homo habilis, Omo offers a unique opportunity to test hypotheses concerning hominid evolution, especially regarding the phylogenetic status of A. africanus. Comparisons of mean cheek teeth breadths disclosed the significant (P less than or equal to 0.05) differences between the Omo sample and the Laetoli/Hadar fossils (P4, M2, and M3), the Homo fossils (P3, P4, M1, M2, and M1), and A. africanus (M3). Of the several possible interpretations of these data, it appears that the high degree of similarity between the Omo sample and the South African gracile australopithecine material warrants considering the two as geographical variants of A. africanus. The geographic, chronologic, and metric attributes of the Omo sample argue for its lineal affinity with A. afarensis and Homo. In conclusion, a consideration of hominid postcanine dental metrics provides no basis for removing A. africanus from the ancestry of the Homo lineage.     

Gluteus maximus muscle function and the origin of hominid bipedality

Bipedality not only frees the hands for tool use but also enhances tool use by allowing use of the trunk for leverage in applying force and thus imparting greater final velocity to tools. Since the weight and acceleration of the trunk and forelimbs on the hindlimbs must be counteracted by muscles such as m. gluteus maximus that control pelvic and trunk movements, it is suggested that the large size of the cranial portion of the human gluteus maximus muscle and its unique attachment to the dorsal ilium (which is apparent in the Makapan australopithecine ilium) may have contributed to the effectiveness with which trunk movement was exploited in early hominid foraging activities. To test this hypothesis, the cranial portions of both right and left muscles were investigated in six human subjects with electromyography during throwing, clubbing, digging, and lifting. The muscles were found to be significantly recruited when the trunk is used in throwing and clubbing, initiating rotation of the pelvis and braking it as trunk rotation ceases and the forelimb accelerates. They stabilize the pelvis during digging and exhibit marked and prolonged activity when the trunk is maintained in partial flexion during lifting of heavy objects.     

Effects of size and locomotor adaptations on the hominid pelvis: evaluation of australopithecine bipedality with a new multivariate method

Three pelves and eight innominate bones belonging to the fossil species, Australopithecus africanus, Australopithecus robustus, Homo erectus, and Homo sapiens, have been studied biometrically and compared with those of recent humans and apes. A new method of logarithmic factorial analysis suppresses both the size effects and the size reference on pelvic proportions. In combination with principal component analysis it allows specializations to be dissociated from allometrical variations. Some morphological differences on the hominid pelvis prove to be mainly allometric. However, the pelvic morphology of australopithecines is clearly differentiated from that of the genus Homo (including H. erectus, OH 28, KNMER 3227). A. africanus (Sts 14, MLD 7, AL 288) is nearer the humans than is A. robustus (SK 50, SK 3155), which appears to be more specialized in the australopithecine lineage. The pelvic morphology of A. africanus, as integrated with the articular pelvic-femoral link, appears to be biometrically equivalent to that of humans.     

Several cranial indices and their relevance to fossil man

An evaluation of three cranial indices proposed by Le Gros Clark ('50) is presented in light of hominid fossil material not previously considered. The data analysis suggests that the supra-orbital height index is an important contribution to cranial analysis and elaborations of it may prove valuable. From the results, the authors suggest that the two australopithecine types be separated on the genus level, i.e., Australopithecus africanus and Paranthropus robustus. It is also proposed that Skhūl V be considered Homo sapiens sapiens. The authors generally conclude that a more dynamic approach in primate cranial analysis is needed. This approach should incorporate non-morphological variables.     

New australopithecine endocast,SK 1585, from Swartkrans,South Africa

The new SK 1585 endocast, found by Dr. Brain at Swartkrans, 1966, is that of a robust australopithecine, matching the endocast of the Olduvai Hominid 5 in volume, and being almost identical to it in morphology. Aside from Olduvai Hominid 5 it is the only robust australopithecine endocast complete enough to permit easy reconstruction, as only a small portion of the frontal lobe is missing. While the gyral and sulcal patterns are not clear, there are a number of features indicating that the brain is not that of a pongid, but that is has been reorganized to a hominid pattern, particularly the occipital, parietal, and temporal lobes.     

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.     

Bipedality in chimpanzee (Pan troglodytes) and bonobo (Pan paniscus): testing hypotheses on the evolution of bipedalism

A host of ecological, anatomical, and physiological selective pressures are hypothesized to have played a role in the evolution of hominid bipedalism. A referential model, based on the chimpanzee (Pan troglodytes) and bonobo (Pan paniscus), was used to test through experimental manipulation four hypotheses on the evolution of hominid bipedalism. The introduction of food piles (Carry hypothesis) increased locomotor bipedality in both species. Neither the introduction of branches (Display hypothesis) nor the construction of visual barriers (Vigilance hypothesis) altered bipedality in either species. Introduction of raised foraging structures (Forage hypothesis) increased postural bipedality in chimpanzees. These experimental manipulations provided support for carrying of portable objects and foraging on elevated food-items as plausible mechanisms that shaped bipedalism in hominids.     

Mandibular postcanine dentition from the Shungura Formation,Ethiopia: Crown morphology,taxonomic allocations,and Plio-Pleistocene hominid evolution

Over 200 hominid specimens were recovered by the International Omo Expedition of 1967–1976. Despite the fragmentary nature of this primarily dental collection, these hominid remains represent a major body of evidence about hominid evolution in eastern Africa during the 2–3 myr time period. Our analysis of the Omo dental collection is based on a large comparative sample of 375 quantifiable mandibular postcanine teeth of A. afarensis, A. africanus, A. aethiopicus, A. boisei, A. robustus, and early Homo. A total of 48 isolated mandibular premolars and molars of the Omo collection spanning the 2–3 myr time period is sufficiently preserved to allow reliable serial allocations and intertaxon comparisons and is the object of study in this paper. We present taxonomic identifications of these teeth and seven other mandibular specimens preserving tooth crowns. Metric analyses of this study include cusp area and crown shape variables taken on occlusal view diagrams. Nonmetric analyses were based on simultaneous observations of all relevant material to ensure accuracy of categorical evaluations. First, a combined metric and morphological evaluation was conducted to allocate each Omo tooth to either robust or nonrobust categories. Further taxonomic affinities were then examined. Our results indicate that nonrobust and robust lineages cooccur by circa 2.7 myr. We consider the Shungura robust specimens from Members C through F to represent A. aethiopicus. A significant phenetic transformation occurs at circa 2.3 myr, with the mosaic emergence of the derived A. boisei morphology across Member G times. Characterization of the East African nonrobust lineage is more difficult because of the comparatively subtle morphological differences seen among the dentitions of A. afarensis, A. africanus, and early Homo. The earlier Members B and C nonrobust specimens are difficult to evaluate and are considered indeterminate to genus or species. Both molars and premolars from Members E through G exhibit phenetic similarities to the early Homo condition and are considered as aff. Homo sp. indet. At present, there is no indication of multiple species in the Omo nonrobust sample at any time horizon. The 2–2.4 myr Omo nonrobust specimens exhibit some similarities to the stated Homo “rudolfensis” condition in size and morphology and are likely to represent the ancestral condition of the genus Homo. The bearing of these results on interpretations of early hominid evolution and diversification is considered. © 1996 Wiley-Liss, Inc.     

From apes to humans: locomotion as a key feature for phylogeny

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).