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Almost a century and a half ago, Charles Darwin in The Descent of Man (1871: 141) highlighted the evolution of bipedalism as one of the key features of the human lineage, freeing the hands for carrying and for using and making tools. But how did it arise? The famous footprints from Laetoli in Tanzania show that hominin ancestors were walking upright by at least 3.63 million years ago. Recent work, however, suggests a much earlier origin for bipedalism, in a Miocene primate ancestor that was still predominantly tree-dwelling. Here Susannah Thorpe, Juliet McClymont and Robin Crompton set out the evidence for that hypothesis and reject the notion that the common ancestor of great apes and humans was a knuckle-walking terrestrial species, as are gorillas and chimpanzees today. The article is followed by a series of comments, rounded off by a reply from the authors., Theories regarding the origins of hominin bipedalism have spent some considerable time 'on the ground' as a result of the knuckle-walking hypothesis, which postulates that our earliest bipedal ancestor evolved [...]

The Asian apes, more than any other, are restricted to an arboreal habitat. They are consequently an important model in the interpretation of the morphological commonalities of the apes, which are locomotor features associated with arboreal living. This paper presents a detailed analysis of orangutan positional behavior for all age-sex categories and during a complete range of behavioral contexts, following standardized positional mode descriptions proposed by Hunt et al. ([1996] Primates 37:363-387). This paper shows that orangutan positional behavior is highly complex, representing a diverse spectrum of positional modes. Overall, all orthograde and pronograde suspensory postures are exhibited less frequently in the present study than previously reported. Orthograde suspensory locomotion is also exhibited less often, whereas pronograde and orthograde compressive locomotor modes are observed more frequently. Given the complexity of orangutan positional behavior demonstrated by this study, it is likely that differences in positional behavior between studies reflect differences in the interplay between the complex array of variables, which were shown to influence orangutan positional behavior (Thorpe and Crompton [2005] Am. J. Phys. Anthropol. 127:58-78). With the exception of pronograde suspensory posture and locomotion, orangutan positional behavior is similar to that of the African apes, and in particular, lowland gorillas. This study suggests that it is orthogrady in general, rather than forelimb suspend specifically, that characterizes the positional behavior of hominoids. KEY WORDS Pongo pygmaeus; posture; orthograde clamber; forelimb suspend

The large body mass and exclusively arboreal lifestyle of Sumatran orangutans identify them as a key species in understanding the dynamic between primates and their environment. Increased knowledge of primate locomotor ecology, coupled with recent developments in the standardization of positional mode classifications (Hunt et al. [1996] Primates 37:363-387), opened the way for sophisticated multivariate statistical approaches, clarifying complex associations between multiple influences on locomotion. In this study we present a log-linear modelling approach used to identify key associations between orangutan locomotion, canopy level, support use, and contextual behavior. Log-linear modelling is particularly appropriate because it is designed for categorical data, provides a systematic method for testing alternative hypotheses regarding interactions between variables, and allows interactions to be ranked numerically in terms of relative importance. Support diameter and type were found to have the strongest associations with locomotor repertoire, suggesting that orangutans have evolved distinct locomotor modes to solve a variety of complex habitat problems. However, height in the canopy and contextual behavior do not directly influence locomotion: instead, their effect is modified by support type and support diameter, respectively. Contrary to classic predictions, age-sex category has only limited influence on orangutan support use and locomotion, perhaps reflecting the presence of arboreal pathways which individuals of all age-sex categories follow. Effects are primarily related to a tendency for adult, parous females to adopt a more cautious approach to locomotion than adult males and immature subjects. KEY WORDS primate; locomotion; support use

In their comment (above), Winder and colleagues refine their original Hypothesis (Winder et al. 2013) to focus explicitly on the transition from arboreal to terrestrial orthogrady. They indicate that they [...]

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-section area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands of the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally. Am J Phys Anthropol 110:179-199, 1999.