Kristian J. Carlson

Australopithecus sediba: A New Species of Homo-Like Australopith from South Africa

From Australopithecus to Homo Our genus Homo is thought to have evolved a little more than 2 million years ago from the earlier hominid Australopithecus. But there are few fossils that provide detailed information on this transition. Berger et al. (p. 195; see the cover) now describe two partial skeletons, including most of the skull, pelvis, and ankle, of a new species of Australopithecus that are informative. The skeletons were found in a cave in South Africa encased in sediments dated by Dirks et al. (p. 205) to about 1.8 to 1.9 million years ago. The fossils share many derived features with the earliest Homo species, including in its pelvis and smaller teeth, and imply that the transition to Homo was in stages. A new species of Australopithecus, about 1.9 million years old, shows many derived features with Homo, helping to reveal its evolution. Despite a rich African Plio-Pleistocene hominin fossil record, the ancestry of Homo and its relation to earlier australopithecines remain unresolved. Here we report on two partial skeletons with an age of 1.95 to 1.78 million years. The fossils were encased in cave deposits at the Malapa site in South Africa. The skeletons were found close together and are directly associated with craniodental remains. Together they represent a new species of Australopithecus that is probably descended from Australopithecus africanus. Combined craniodental and postcranial evidence demonstrates that this new species shares more derived features with early Homo than any other australopith species and thus might help reveal the ancestor of that genus.

The Foot and Ankle of Australopithecus sediba

Australopithecus sediba had a human-like ankle and arch but an ape-like heel and tibia, implying that while bipedal, this species was also adept at climbing trees. A well-preserved and articulated partial foot and ankle of Australopithecus sediba, including an associated complete adult distal tibia, talus, and calcaneus, have been discovered at the Malapa site, South Africa, and reported in direct association with the female paratype Malapa Hominin 2. These fossils reveal a mosaic of primitive and derived features that are distinct from those seen in other hominins. The ankle (talocrural) joint is mostly humanlike in form and inferred function, and there is some evidence for a humanlike arch and Achilles tendon. However, Au. sediba is apelike in possessing a more gracile calcaneal body and a more robust medial malleolus than expected. These observations suggest, if present models of foot function are correct, that Au. sediba may have practiced a unique form of bipedalism and some degree of arboreality. Given the combination of features in the Au. sediba foot, as well as comparisons between Au. sediba and older hominins, homoplasy is implied in the acquisition of bipedal adaptations in the hominin foot.

The Endocast of MH1, Australopithecus sediba

The brain endocast of Australopithecus sediba shows that despite retaining a small brain size, some reorganization of the frontal lobe had commenced, hinting at the later neural development seen in Homo. The virtual endocast of MH1 (Australopithecus sediba), obtained from high-quality synchrotron scanning, reveals generally australopith-like convolutional patterns on the frontal lobes but also some foreshadowing of features of the human frontal lobes, such as posterior repositioning of the olfactory bulbs. Principal component analysis of orbitofrontal dimensions on australopith endocasts (MH1, Sts 5, and Sts 60) indicates that among these, MH1 orbitofrontal shape and organization align most closely with human endocasts. These results are consistent with gradual neural reorganization of the orbitofrontal region in the transition from Australopithecus to Homo, but given the small volume of the MH1 endocast, they are not consistent with gradual brain enlargement before the transition.

A Partial Pelvis of Australopithecus sediba

Although it had a small brain and skull, Australopithecus sediba shows some human-like features in its reconstructed pelvis. The fossil record of the hominin pelvis reflects important evolutionary changes in locomotion and parturition. The partial pelves of two individuals of Australopithecus sediba were reconstructed from previously reported finds and new material. These remains share some features with australopiths, such as large biacetabular diameter, small sacral and coxal joints, and long pubic rami. The specimens also share derived features with Homo, including more vertically oriented and sigmoid-shaped iliac blades, greater robusticity of the iliac body, sinusoidal anterior iliac borders, shortened ischia, and more superiorly oriented pubic rami. These derived features appear in a species with a small adult brain size, suggesting that the birthing of larger-brained babies was not driving the evolution of the pelvis at this time.

The Lower Limb and Mechanics of Walking in Australopithecus sediba

The discovery of a relatively complete Australopithecus sediba adult female skeleton permits a detailed locomotor analysis in which joint systems can be integrated to form a comprehensive picture of gait kinematics in this late australopith. Here we describe the lower limb anatomy of Au. sediba and hypothesize that this species walked with a fully extended leg and with an inverted foot during the swing phase of bipedal walking. Initial contact of the lateral foot with the ground resulted in a large pronatory torque around the joints of the foot that caused extreme medial weight transfer (hyperpronation) into the toe-off phase of the gait cycle (late pronation). These bipedal mechanics are different from those often reconstructed for other australopiths and suggest that there may have been several forms of bipedalism during the Plio-Pleistocene.

Is Bone's Response to Mechanical Signals Dominated by Gravitational Loading?

During locomotion and exercise, bone is subjected to forces induced by gravitational loading and muscle loading. The inherent link between these modes of loading has confounded emergence of either one as the principal anabolic or anticatabolic signal in bone. A paradigm has emerged in the literature stipulating that muscle loading is the larger of the two, and therefore, bone morphology is predominantly determined by muscle loads. In spite of the intuitive appeal of a muscle-bone unit tuned to the magnitude of contractile forces, little evidence exists for the relatively few, large-magnitude muscle contractions arising during daily activities to dominate the mechanosensory input of bone. Moreover, a review of the literature raises several inconsistencies in this paradigm and indicates that the alternative--gravitational loading--can have a significant role in determining bone mass and morphology. Certainly, the relative contribution of each type of loading will depend on the specific activity, the location of the bone within the skeleton, and whether the bone is weight-bearing or not. Most likely, a more comprehensive paradigm for explaining sensitivity of bone to loading will have to include not only large-magnitude gravitational and muscle loads, but also other factors such as high-frequency, low-magnitude signals generated by the muscles during postural adjustments.

The Upper Limb of Australopithecus sediba

The evolution of the human upper limb involved a change in function from its use for both locomotion and prehension (as in apes) to a predominantly prehensile and manipulative role. Well-preserved forelimb remains of 1.98-million-year-old Australopithecus sediba from Malapa, South Africa, contribute to our understanding of this evolutionary transition. Whereas other aspects of their postcranial anatomy evince mosaic combinations of primitive (australopith-like) and derived (Homo-like) features, the upper limbs (excluding the hand and wrist) of the Malapa hominins are predominantly primitive and suggest the retention of substantial climbing and suspensory ability. The use of the forelimb primarily for prehension and manipulation appears to arise later, likely with the emergence of Homo erectus.

Increased non-linear locomotion alters diaphyseal bone shape.

SUMMARY Comparative studies of vertebrate morphology that link habitual locomotor activities to bone structural properties are often limited by confounding factors such as genetic variability between groups. Experimental assessment of bones adaptive response to altered activity patterns typically involves superimposing exercise onto a normal locomotor repertoire, making a distinction between qualitative changes to locomotor repertoires and quantitative increases in activity level difficult. Here, we directly tested the hypothesis that an increase in turning activity, without the application of exercise per se, will alter femoral cross-sectional shape. Thirty day-old female BALB/cByJ mice (n=10 per group) were single-housed for 8 weeks in custom-designed cages that either accentuated linear or turning locomotion or allowed subjects to freely roam standard cages. Consistent with a lack of difference in physical activity levels between groups, there were no significant differences in body mass, femoral length, midshaft cortical area, and individual measures of mediolateral (ML) and anteroposterior (AP) bending rigidity. However, the ratio of ML to AP diaphyseal rigidity, an indicator of cross-sectional shape, was significantly greater (P<0.05) in turning subjects than in linear or control subjects. Considering that across all groups mice were genetically identical and had equivalent levels of bone quantity and physical activity, differences in femoral shape were attributed to qualitative differences in locomotor patterns (i.e. specific locomotor modes). These data indicate that increased turning can alter distribution of bone mass in the femoral diaphysis, and that turning should be considered in efforts to understand form–function relationships in vertebrates.

Recent origin of low trabecular bone density in modern humans

Significance The human skeleton is unique in having low trabecular density representing a lightly built human body form. However, it remains unknown when during human evolution this unique characteristic first appeared. To our knowledge, this study is the first to examine trabecular bone density throughout the skeleton of fossil hominins spanning several million years. The results show that trabecular density remained high throughout human evolution until it decreased significantly in recent modern humans, suggesting a possible link between changes in our skeleton and increased sedentism. Humans are unique, compared with our closest living relatives (chimpanzees) and early fossil hominins, in having an enlarged body size and lower limb joint surfaces in combination with a relatively gracile skeleton (i.e., lower bone mass for our body size). Some analyses have observed that in at least a few anatomical regions modern humans today appear to have relatively low trabecular density, but little is known about how that density varies throughout the human skeleton and across species or how and when the present trabecular patterns emerged over the course of human evolution. Here, we test the hypotheses that (i) recent modern humans have low trabecular density throughout the upper and lower limbs compared with other primate taxa and (ii) the reduction in trabecular density first occurred in early Homo erectus, consistent with the shift toward a modern human locomotor anatomy, or more recently in concert with diaphyseal gracilization in Holocene humans. We used peripheral quantitative CT and microtomography to measure trabecular bone of limb epiphyses (long bone articular ends) in modern humans and chimpanzees and in fossil hominins attributed to Australopithecus africanus, Paranthropus robustus/early Homo from Swartkrans, Homo neanderthalensis, and early Homo sapiens. Results show that only recent modern humans have low trabecular density throughout the limb joints. Extinct hominins, including pre-Holocene Homo sapiens, retain the high levels seen in nonhuman primates. Thus, the low trabecular density of the recent modern human skeleton evolved late in our evolutionary history, potentially resulting from increased sedentism and reliance on technological and cultural innovations.

Muscle architecture of the common chimpanzee ( Pan troglodytes ): perspectives for investigating chimpanzee behavior

Thorpe et al. (Am J Phys Anthropol 110:179–199, 1999) quantified chimpanzee (Pan troglodytes) muscle architecture and joint moment arms to determine whether they functionally compensated for structural differences between chimpanzees and humans. They observed enough distinction to conclude that musculoskeletal properties were not compensatory and suggested that chimpanzees and humans do not exhibit dynamically similar movements. These investigators based their assessment on unilateral limb musculatures from three male chimpanzees, of which they called one non-adult representative. Factors such as age, sex, and behavioral lateralization may be responsible for variation in chimpanzee muscle architecture, but this is presently unknown. While the full extent of variation in chimpanzee muscle architecture due to such factors cannot be evaluated with data presently available, the present study expands the chimpanzee dataset and provides a preliminary glimpse of the potential relevance of these factors. Thirty-seven forelimb and 36 hind limb muscles were assessed in two chimpanzee cadavers: one unilaterally (right limbs), and one bilaterally. Mass, fiber length, and physiological cross-sectional area (PCSA) are reported for individual muscles and muscle groups. The musculature of an adult female is more similar in architectural patterns to a young male chimpanzee than to humans, particularly when comparing muscle groups. Age- and sex-related intraspecific differences do not obscure chimpanzee-human interspecific differences. Side asymmetry in one chimpanzee, despite consistent forelimb directional asymmetry, also does not exceed the magnitude of chimpanzee-human differences. Left forelimb muscles, on average, usually had higher masses and longer fiber lengths than right, while right forelimb muscles, on average, usually had greater PCSAs than left. Most muscle groups from the left forelimb exhibited greater masses than right groups, but group asymmetry was significant only for the manual digital muscles. The hind limb exhibited less asymmetry than the forelimb in most comparisons. Examination of additional chimpanzees would clarify the full range of inter- and intra-individual variation.