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The StW 573 skeleton of Australopithecus prometheus from Sterkfontein Member 2 is some 93% complete and thus by far the most complete member of that genus yet found. Firmly dated at 3.67 Ma, it is one of the earliest specimens of its genus. A crucial aspect of interpretation of locomotor behaviour from fossil remains is an understanding of the palaeoenvironment in which the individual lived and the manner in which it would have used it. While the value of this ecomorphological approach is largely accepted, it has not been widely used as a stable framework on which to build evolutionary biomechanical interpretations. Here, we collate the available evidence on StW 573’s anatomy in order, as far as currently possible, to reconstruct what might have been this individual’s realized and potential niche. We explore the concept of a common Australopithecus “bauplan” by comparing the morphology and ecological context of StW 573 to that of paenocontemporaneous australopiths including Australopithecus anamensis and KSD-VP-1/1 Australopithecus afarensis. Each was probably substantially arboreal and woodland-dwelling, relying substantially on arboreal resources. We use a hypothesis-driven approach, tested by: virtual experiments, in the case of extinct species; biomechanical analyses of the locomotor behaviour of living great ape species; and analogical experiments with human subjects. From these, we conclude that the habitual locomotor mode of all australopiths was upright bipedalism, whether on the ground or on branches. Some later australopiths such as Australopithecus sediba undoubtedly became more terrestrial, allowing sacrifice of arboreal stability in favour of manual dexterity. Indeed, modern humans retain arboreal climbing skills but have further sacrificed arboreal effectiveness for enhanced ability to sustain striding terrestrial bipedalism over much greater distances. We compare StW 573’s locomotor adaptations to those of living great apes and protohominins, and agree with those earlier observers who suggest that the common panin-hominin last common ancestor was postcranially more like Gorilla than Pan.

Functional morphology of the atlas reflects multiple aspects of an organism’s biology. More specifically, its shape indicates patterns of head mobility, while the size of its vascular foramina reflects blood flow to the brain. Anatomy and function of the early hominin atlas, and thus, its evolutionary history, are poorly documented because of a paucity of fossilized material. Meticulous excavation, cleaning and high-resolution micro-CT scanning of the StW 573 (‘Little Foot’) skull has revealed the most complete early hominin atlas yet found, having been cemented by breccia in its displaced and flipped over position on the cranial base anterolateral to the foramen magnum. Description and landmark-free morphometric analyses of the StW 573 atlas, along with other less complete hominin atlases from Sterkfontein (StW 679) and Hadar (AL 333-83), confirm the presence of an arboreal component in the positional repertoire of Australopithecus. Finally, assessment of the cross-sectional areas of the transverse foramina of the atlas and the left carotid canal in StW 573 further suggests there may have been lower metabolic costs for cerebral tissues in this hominin than have been attributed to extant humans and may support the idea that blood perfusion of these tissues increased over the course of hominin evolution.

The ca. 3.67 Ma adult skeleton known as ‘Little Foot’ (StW 573), recovered from Sterkfontein Member 2 breccia in the Silberberg Grotto, is remarkable for its morphology and completeness. Preservation of clavicles and scapulae, including essentially complete right-side elements, offers opportunities to assess morphological and functional aspects of a nearly complete Australopithecus pectoral girdle. Here we describe the StW 573 pectoral girdle and offer quantitative comparisons to those of extant hominoids and selected homininans. The StW 573 pectoral girdle combines features intermediate between those of humans and other apes: a long and curved clavicle, suggesting a relatively dorsally positioned scapula; an enlarged and uniquely proportioned supraspinous fossa; a relatively cranially oriented glenoid fossa; and ape-like reinforcement of the axillary margin by a stout ventral bar. StW 573 scapulae are as follows: smaller than those of some homininans (i.e., KSD-VP-1/1 and KNM-ER 47000A), larger than others (i.e., A.L. 288-1, Sts 7, and MH2), and most similar in size to another australopith from Sterkfontein, StW 431. Moreover, StW 573 and StW 431 exhibit similar structural features along their axillary margins and inferior angles. As the StW 573 pectoral girdle (e.g., scapular configuration) has a greater affinity to that of apes—Gorilla in particular—rather than modern humans, we suggest that the StW 573 morphological pattern appears to reflect adaptations to arboreal behaviors, especially those with the hand positioned above the head, more than human-like manipulatory capabilities. When compared with less complete pectoral girdles from middle/late Miocene apes and that of the penecontemporaneous KSD-VP-1/1 (Australopithecus afarensis), and mindful of consensus views on the adaptiveness of arboreal positional behaviors soliciting abducted glenohumeral joints in early Pliocene taxa, we propose that the StW 573 pectoral girdle is a reasonable model for hypothesizing pectoral girdle configuration of the crown hominin last common ancestor.

Due to difficulty of obtaining accurate quantitative data on foot muscles, relatively little has been done to study foot muscle function in non-human apes. Gorilla feet are known to be similar in bony proportions and mechanics to those of humans, hence are key to understanding human foot evolution and its ecological context. We present the first 3D musculoskeletal computer model of a western lowland gorilla foot, giving muscle torques about the tarsometatarsal, metatarsophalangeal and interphalangeal joints of digits 2–5. Peak flexor torque around the fifth metatarsophalangeal joint occurs at a highly flexed position, suggesting an ability to maintain flexed postures around lateral metatarsophalangeal joints, useful for grasping vertical supports. For distal interphalangeal joints, flexor torques peaked the more medial the digit at relatively flexed postures. We report, for the first time, interossei acting upon proximal and distal interphalangeal joints. All these facilitate maintenance of flexed positions around distal interphalangeal joints, likely used for grasping of small supports/objects. Humans lack these features, suggesting that semi-arboreal early hominins made less use of the peripheral canopy than gorillines. Information here could be used in gorilla enclosure design to encourage wild-type locomotor repertoires in captivity.

International audience; Because of its exceptional degree of preservation and its geological age of ∼3.67 Ma, StW 573 makes an invaluable contribution to our understanding of early hominin evolution and paleobiology. The morphology of the bony labyrinth has the potential to provide information about extinct primate taxonomic diversity, phylogenetic relationships and locomotor behaviour. In this context, we virtually reconstruct and comparatively assess the bony labyrinth morphology in StW 573. As comparative material, we investigate 17 southern African hominin specimens from Sterkfontein, Swartkrans and Makapansgat (plus published data from two specimens from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant chimpanzee specimens. We apply a landmark-based geometric morphometric method for quantitatively assessing labyrinthine morphology. Morphology of the inner ear in StW 573 most closely resembles that of another Australopithecus individual from Sterkfontein, StW 578, recovered from the Jacovec Cavern. Within the limits of our sample, we observe a certain degree of morphological variation in the Australopithecus assemblage of Sterkfontein Member 4. Cochlear morphology in StW 573 is similar to that of other Australopithecus as well as to Paranthropus specimens included in this study, but it is substantially different from early Homo. Interestingly, the configuration of semicircular canals in Paranthropus specimens from Swartkrans differs from other fossil hominins, including StW 573. Given the role of the cochlea in the sensory-driven interactions with the surrounding environment, our results offer new perspectives for interpreting early hominin behaviour and ecology. Finally, our study provides additional evidence for discussing the phylogenetic polarity of labyrinthine traits in southern African hominins.

It is traditionally held that early hominins of the genus Australopithecus had a foot transitional in function between that of the other great apes and our own but that the appearance of genus Homo was marked by evolution of an essentially biomechanically modern foot, as well as modern body proportions. Here, we report the application of whole foot, pixel-wise topological statistical analysis, to compare four populations of footprints from across evolutionary time: Australopithecus at Laetoli (3.66 Ma, Tanzania), early African Homo from Ileret (1.5 Ma, Kenya) and recent modern (presumptively habitually barefoot) pastoralist Homo sapiens from Namibia (Holocene), with footprints from modern Western humans. Contrary to some previous analyses, we find that only limited areas of the footprints show any statistically significant difference in footprint depth (used here as an analogy for plantar pressure). A need for this comparison was highlighted by recent studies using the same statistical approach, to examine variability in the distribution of foot pressure in modern Western humans. This study revealed very high intra-variability (mean square error) step-to-step in over 500 steps. This result exemplifies the fundamental movement characteristic of dynamic biological systems, whereby regardless of the repetition in motor patterns for stepping, and even when constrained by experimental conditions, each step is unique or non-repetitive; hence, repetition without repetition. Thus, the small sample sizes predominant in the fossil and ichnofossil record do not reveal the fundamental neurobiological driver of locomotion (variability), essentially limiting our ability to make reliable interpretations which might be extrapolated to interpret hominin foot function at a population level. However, our need for conservatism in our conclusions does not equate with a conclusion that there has been functional stasis in the evolution of the hominin foot.

Background Recent work using large datasets (>500 records per subject) has demonstrated seemingly high levels of step-to-step variation in peak plantar pressure within human individuals during walking. One intuitive consequence of this variation is that smaller sample sizes (e.g., 10 steps per subject) may be quantitatively and qualitatively inaccurate and fail to capture the variance in plantar pressure of individuals seen in larger data sets. However, this remains quantitatively unexplored reflecting a lack of detailed investigation of intra-subject sample size effects in plantar pressure analysis. Methods Here we explore the sensitivity of various plantar pressure metrics to intra-subject sample size (number of steps per subject) using a random subsampling analysis. We randomly and incrementally subsample large data sets (>500 steps per subject) to compare variability in three metric types at sample sizes of 5–400 records: (1) overall whole-record mean and maximum pressure; (2) single-pixel values from five locations across the foot; and (3) the sum of pixel-level variability (measured by mean square error, MSE) from the whole plantar surface. Results Our results indicate that the central tendency of whole-record mean and maximum pressure within and across subjects show only minor sensitivity to sample size >200 steps. However, 400 steps. The range in pixel-level MSE at low subsamples (500 pressure records per subject. Overall, therefore, we demonstrate a high probability that the very small sample sizes (n

Background Most research investigating the connection between walking and visual behaviour has assessed only eye movements (not head orientation) in respect to locomotion over smooth surfaces in a laboratory. This is unlikely to reflect gaze changes found over the complex surfaces experienced in the real world, especially given that eye and head movements have rarely been assessed simultaneously. Research question How does gaze (eye and head) angle and gait speed change when walking over surfaces of different complexity? Methods In this exploratory study, we used a mobile eye tracker to monitor eye movements and inertia measurement unit sensors (IMUs) to measure head angle whilst subjects (n = 11) walked over surfaces with different complexities both indoors and outdoors. Gait speed was recorded from ankle IMUs. Results Overall, mean gaze angle was lowest over the most complex surface and this surface also elicited the slowest mean gait speed. The head contributed increasingly to the lowering of gaze with increased surface complexity. Less complex surfaces showed no significant difference between gaze and gait behaviour. Significance This study supports previous research showing that increased surface complexity is an important factor in determining gaze and gait behaviour. Moreover, it provides the novel finding that head movements provide important contributions to gaze location. Our future research aims are to further assess the role of the head in determining gaze location during locomotion across a greater range of complex surfaces to determine the key surface characteristics that influence gaze during gait.

Background Walking surfaces vary in complexity and are known to affect stability and fall risk whilst walking. However, existing studies define surfaces through descriptions only. Objective This study used a multimethod approach to measure surface complexity in order to try to characterise surfaces with respect to locomotor stability. Methods We assessed how physical measurements of walking surface complexity compared to participant's perceptual ratings of the effect of complexity on stability. Physical measurements included local slope measures from the surfaces themselves and shape complexity measured using generated surface models. Perceptual measurements assessed participants' perceived stability and surface roughness using Likert scales. We then determined whether these measurements were indicative of changes to stability as assessed by behavioural changes including eye angle, head pitch angle, muscle coactivation, walking speed and walking smoothness. Results Physical and perceptual measures were highly correlated, with more complex surfaces being perceived as more challenging to stability. Furthermore, complex surfaces, as defined from both these measurements, were associated with lowered head pitch, increased muscle coactivation and reduced walking smoothness. Significance Our findings show that walking surfaces defined as complex, based on physical measurements, are perceived as more challenging to our stability. Furthermore, certain behavioural measures relate better to these perceptual and physical measures than others. Crucially, for the first time this study defined walking surfaces objectively rather than just based on subjective descriptions. This approach could enable future researchers to compare results across walking surface studies. Moreover, perceptual measurements, which can be collected easily and efficiently, could be used as a proxy for estimating behavioural responses to different surfaces. This could be particularly valuable when determining risk of instability when walking for individuals with compromised stability.

International audience; The Sterkfontein Caves is currently the world's richest Australopithecus-bearing site. Included in Sterkfontein's hominin assemblage is StW 573 (‘Little Foot’), a near-complete Australopithecus skeleton discovered in Member 2 in the Silberberg Grotto. Because of its importance to the fossil hominin record, the geological age of StW 573 has been the subject of significant debate. Three main hypotheses have been proposed regarding the formation and age of Member 2 and by association StW 573. The first proposes that Member 2 (as originally defined in the type section in the Silberberg Grotto) started to accumulate at around 2.58 Ma and that the unit is contained within the Silberberg Grotto. The second proposes that Member 2 started forming before 3.67 ± 0.16 Ma and that the deposit extends into the Milner Hall and close to the base of the cave system. The third proposes a ‘two-stage burial scenario’, in which some sediments and StW 573 represent a secondary and mixed-age accumulation reworked from a higher cave. The stratigraphic and sedimentological implications of these hypotheses are tested here through the application of a multiscale investigation of Member 2, with reference to the taphonomy of the StW 573 skeleton. The complete infilling sequence of Member 2 is described across all exposures of the deposit in the Silberberg Grotto and into the Milner Hall. Sediments are generally stratified and conformably deposited in a sequence of silty sands eroded from well-developed lateritic soils on the landscape surface. Voids, clasts and bioclasts are organized consistently across and through Member 2 conforming with the underlying deposit geometry, indicating gradual deposit accretion with no distinct collapse facies evident and only localized intra-unit postdepositional modification. The stratigraphy and sedimentology of Member 2 support a simple single-stage accumulation process of Member 2 and a primary association between the sediments of Member 2 and the StW 573 ‘Little Foot’ skeleton.

In the early 20th century the dominant paradigm for the ecological context of the origins of human bipedalism was arboreal suspension. In the 1960s, however, with recognition of the close genetic relationship of humans, chimpanzees and bonobos, and with the first field studies of mountain gorillas and common chimpanzees, it was assumed that locomotion similar to that of common chimpanzees and mountain gorillas, which appeared to be dominated by terrestrial knuckle-walking, must have given rise to human bipedality. This paradigm has been popular, if not universally dominant, until very recently. However, evidence that neither the knuckle-walking or vertical climbing of these apes is mechanically similar to human bipedalism, as well as the hand-assisted bipedality and orthograde clambering of orang-utans, has cast doubt on this paradigm. It now appears that the dominance of terrestrial knuckle-walking in mountain gorillas is an artefact seen only in the extremes of their range, and that both mountain and lowland gorillas have a generalized orthogrady similar to that seen in orang-utans. These data, together with evidence for continued arboreal competence in humans, mesh well with an increasing weight of fossil evidence suggesting that a mix of orang-utan and gorilla-like arboreal locomotion and upright terrestrial bipedalism characterized most australopiths. The late split date of the panins, corresponding to dates for separation of Homo and Australopithecus, leads to the speculation that competition with chimpanzees, as appears to exist today with gorillas, may have driven ecological changes in hominins and perhaps cladogenesis. However, selection for ecological plasticity and morphological conservatism is a core characteristic of Hominidae as a whole, including Hominini.

Background Peripheral vision often deteriorates with age, disrupting our ability to maintain normal locomotion. Laboratory based studies have shown that lower visual field loss, in particular, is associated with changes in gaze and gait behaviour whilst walking and this, in turn, increases the risk of falling in the elderly. Separately, gaze and gait behaviours change and fall risk increases when walking over complex surfaces. It seems probable, but has not yet been established, that these challenges to stability interact. Research question How does loss of the lower visual field affect gaze and gait behaviour whilst walking on a variety of complex surfaces outside of the laboratory? Specifically, is there a synergistic interaction between the effects on behaviour of blocking the lower visual field and increased surface complexity? Methods We compared how full vision versus simulated lower visual field loss affected a diverse range of behavioural measures (head pitch angle, eye angle, muscle coactivation, gait speed and walking smoothness as measured by harmonic ratios) in young participants. Participants walked over a range of surfaces of different complexity, including pavements, grass, steps and pebbles. Results In both full vision and blocked lower visual field conditions, surface complexity influenced gaze and gait behaviour. For example, more complex surfaces were shown to be associated with lowered head pitch angles, increased leg muscle coactivation, reduced gait speed and decreased walking smoothness. Relative to full vision, blocking the lower visual field caused a lowering of head pitch, especially for more complex surfaces. However, crucially, muscle coactivation, gait speed and walking smoothness did not show a significant change between full vision and blocked lower visual field conditions. Finally, head pitch angle, muscle coactivation, gait speed and walking smoothness were all correlated highly with each other. Significance Our study showed that blocking the lower visual field did not significantly change muscle coactivation, gait speed or walking smoothness. This suggests that young people cope well when walking with a blocked lower visual field, making minimal behavioural changes. Surface complexity had a greater effect on gaze and gait behaviour than blocking the lower visual field. Finally, head pitch angle was the only measure that showed a significant synergistic interaction between surface complexity and blocking the lower visual field. Together our results indicate that, first, a range of changes occur across the body when people walk over more complex surfaces and, second, that a relatively simple behavioural change (to gaze) suffices to maintain normal gait when the lower visual field is blocked, even in more challenging environments. Future research should assess whether young people cope as effectively when several impairments are simulated, representative of the comorbidities found with age.

The Sterkfontein Caves has an 80 year history of yielding remarkable evidence of hominin evolution and is currently the world’s richestAustralopithecus-beafingsite. Included in Sterkfontein’s hominin assemblage is StW 573 (‘Little Foot’). Discovered in the Member 2 deposit in the Silberberg Grotto, StW 573 represents the most completeAustralopithecusskeleton yet found. Because of its importance to the fossil hominin record, the geological age of Little Foot has been the subject of significant debate. Two main hypotheses have been proposed regarding the formation and age of Member 2 and by association StW 573. The first, proposes that Member 2 formed relatively rapidly, starting to accumulate at around 2.8 million years ago and that the unit is isolated to the Silberberg Grotto - the underlying chambers and passages forming later. The second proposes that Member 2 formed slowly, its accumulation starting before 3.67 million years ago and that the deposit extends into the Milner Hall and close to the base of the cave system. Both assume a primary association between StW 573 and Member 2, although which sediments in the Silberberg Grotto are associated with Member 2 has also been questioned. Recently a third alternative hypothesis questioning the association of StW 573 to Member 2 sediments proposed a ‘two-stage burial scenario’ in which sediments associated with StW 573 represent a secondary and mixed-age deposit reworked from a higher cave. The stratigraphic and sedimentological implications of these hypotheses are tested here through the application of a multiscale investigation of Member 2, with reference to the taphonomy of the Little Foot skeleton. The complete infilling sequence of Member 2 is described and depositional units are tracked across all exposures of the deposit in the Silberberg Grotto and into the Milner Hall. Facies development follows patterns characteristic of colluvially accumulated taluses with 30-40° angles of repose developing coarser distal facies. Sediments are generally stratified and conformably deposited in a sequence of silty sands eroded from well-developed lateritic soils on the landscape surface. Voids, clasts and bioclasts are organized consistently across and through Member 2 according to the underlying deposit geometry, indicating a gradual deposit accretion with no distinct collapse facies evident, no successive debris flow accumulation, and only localized intra-unit post- depositional modification. The stratigraphy and sedimentology of Member 2 supports a simple single-stage accumulation process through which Member 2 partially fills the Silberberg Grotto and extends into the deeper chambers and passages of the Sterkfontein Caves. Through this work we demonstrate at multiple scales the primary association between the sediments of Member 2 and the StW 573 ‘Little Foot’ skeleton.

Due to its completeness, the A.L. 288-1 (Lucy) skeleton has long served as the archetypal bipedal Australopithecus. However, there remains considerable debate about its limb proportions. There are three competing, but not necessarily mutually exclusive, explanations for the high humerofemoral index of A.L. 288-1: (1) a retention of proportions from an Ardipithecus-like most recent common ancestor (MRCA); (2) indication of some degree of climbing ability; (3) allometry. Recent discoveries of other partial skeletons of Australopithecus, such as those of A. sediba (MH1 and MH2) and A. afarensis (KSD-VP-1/1 and DIK-1/1), have provided new opportunities to test hypotheses of early hominin body size and limb proportions. Yet, no early hominin is as complete (>90%), as is the 3.67 Ma Little Foot (StW 573) specimen, from Sterkfontein Member 2. Here, we provide the first descriptions of its upper and lower long limb bones, as well as a comparative context of its limb proportions. As to the latter, we found that StW 573 possesses absolutely longer limb lengths than A.L. 288-1, but both skeletons show similar limb proportions. This finding seems to argue against an allometric explanation for the limb proportions of A.L. 288-1. In fact, our multivariate allometric analysis suggests that limb lengths of Australopithecus, as represented by StW 573 and A.L. 288-1, developed along a significantly different (p < 0.001) allometric scale than that which typifies modern humans and African apes. Our analyses also suggest, as have those of others, that hominin limb evolution occurred in two stages with: (1) a modest increase in lower limb length and a concurrent shortening of the antebrachium between Ardipithecus and Australopithecus, followed by (2) considerable lengthening of the lower limb along with a decrease of both upper limb elements occurring between Australopithecus and Homo sapiens.

The locomotor behaviour of 2 groups of Propithecus verreauxi (Verreaux's sifaka) was studied over an 8-month period in Kirindy Mitea National Park (KMNP), Madagascar. This paper assesses the major characteristics of their locomotion, focusing on the extent that seasonal variation in climate and habitat, and local variation in habitat, is reflected in changes in locomotor behaviour. P. verreauxi is a committed leaper with a strong preference for vertical and angled supports. We found clear between-group differences in support orientation and diameter suggesting local variation in habitat. During the dry season, P. verreauxi utilizes smaller-diameter supports than in the rainy season. While this difference cannot yet be ascribed to any single cause, we discuss the factors which may contribute to this result.

Three‐dimensional musculoskeletal models have become increasingly common for investigating muscle moment arms in studies of vertebrate locomotion. In this study we present the first musculoskeletal model of a western lowland gorilla hind limb. Moment arms of individual muscles around the hip, knee and ankle were compared with previously published data derived from the experimental tendon travel method. Considerable differences were found which we attribute to the different methodologies in this specific case. In this instance, we argue that our 3D model provides more accurate and reliable moment arm data than previously published data on the gorilla because our model incorporates more detailed consideration of the 3D geometry of muscles and the geometric constraints that exist on their lines‐of‐action about limb joints. Our new data have led us to revaluate the previous conclusion that muscle moment arms in the gorilla hind limb are optimised for locomotion with crouched or flexed limb postures. Furthermore, we found that bipedalism and terrestrial quadrupedalism coincided more regularly with higher moment arms and torque around the hip, knee and ankle than did vertical climbing. This indicates that the ability of a gorilla to walk bipedally is not restricted by musculoskeletal adaptations for quadrupedalism and vertical climbing, at least in terms of moment arms and torque about hind limb joints.

There are at present few comparable studies of lemur locomotion in the wild. This has unfortu- nately meant we have little knowledge about locomotor variation, and hence flexibility, with regard to differences in support availability and habitat structure. Here we compare the locomotion of Lepilemur edwardsi at Ankar- afantsika with that of Lepilemur ruficaudatus at Kirindy-Mitea National Park. While data were collected by two individuals, at different times, both studies used the same data collection protocol and are hence highly comparable. Locomotor mode, support diameter and ori- entation, heights, and distances traveled were all col- lected. We find that locomotor specialization, in this case for vertical leaping, has ensured that some support requirements are independent of habitat. For example, both species used vertical supports most often. However, overall support diameter does indicate a certain degree of flexibility, whereby L. ruficaudatus most often used sup- ports � 5 cm in diameter and L. edwardsi >5 cm in diam- eter. Am J Phys Anthropol 000:000-000, 2014. V C 2014

Here we report on a Holocene inter-dune site close to Walvis Bay (Namibia) which contains exceptionally well-preserved children’s footprints. The footprint surface is dated using Optically Stimulated Luminescence (OSL) methods to approximately 1.5 ka. These dates are compared to those obtained at nearby footprint sites and used to verify a model of diachronous footprint surfaces and also add to the archaeological data available for the communities that occupied these near-coastal areas during the Holocene. This model of diachronous footprint surfaces has implications for other soft-sediment footprint sites such as the 1.5 Ma old footprints at Ileret (Kenya). The distribution of both human and animal tracks, is consistent with the passage of small flock of small ungulates (probably sheep/goats) followed by a group of approximately 9 ± 2 individuals (children or young adults). Age estimates from the tracks suggest that some of the individuals may have been as young as five years old. Variation in track topology across this sedimentologically uniform surface is explained in terms of variations in gait and weight/stature of the individual print makers and is used to corroborate a model of footprint morphology developed at a nearby site. The significance of the site within the literature on human footprints lies in the quality of the track preservation, their topological variability despite a potentially uniform substrate, and the small size of the tracks, and therefore the inferred young age of the track-makers. The site provides an emotive insight into the life of the track-makers.

Almost a century and a half ago, Charles Darwin inThe 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 3.65 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.

The feet of apes have a different morphology from those of humans. Until now, it has merely been assumed that the morphology seen in humans must be adaptive for habitual bipedal walking, as the habitual use of bipedal walking is generally regarded as one of the most clear-cut differences between humans and apes. This study asks simply whether human skeletal proportions do actually enhance foot performance during human-like bipedalism, by examining the influence of foot proportions on force, torque and work in the foot joints during simulated bipedal walking. Skeletons of the common chimpanzee, orangutan, gorilla and human were represented by multi-rigid-body models, where the components of the foot make external contact via finite element surfaces. The models were driven by identical joint motion functions collected from experiments on human walking. Simulated contact forces between the ground and the foot were found to be reasonably comparable with measurements made during human walking using pressure- and force-platforms. Joint force, torque and work in the foot were then predicted. Within the limitations of our model, the results show that during simulated human-like bipedal walking, (1) the human and non-human ape (NHA) feet carry similar joint forces, although the distributions of the forces differ; (2) the NHA foot incurs larger joint torques than does the human foot, although the human foot has higher values in the first tarso-metatarsal and metatarso-phalangeal joints, whereas the NHA foot incurs higher values in the lateral digits; and (3) total work in the metatarso-phalangeal joints is lower in the human foot than in the NHA foot. The results indicate that human foot proportions are indeed well suited to performance in normal human walking.

Center of pressure (COP) trajectories summarize the complex mechanical interaction between the foot and a contacted surface. Each trajectory itself is also complex, comprising hundreds of instantaneous vectors over the duration of stance phase. To simplify statistical analysis often a small number of scalars are extracted from each COP trajectory. The purpose of this paper was to demonstrate how a more objective approach to COP analysis can avoid particular sensitivities of scalar extraction analysis. A previously published dataset describing the effects of walking speed on plantar pressure (PP) distributions was re-analyzed. After spatially and temporally normalizing the data, speed effects were assessed using a vector-field paired Hotelling's T-2 test. Results showed that, as walking speed increased, the COP moved increasingly posterior at heel contact, and increasingly laterally and anteriorly between similar to 60 and 85% stance, in agreement with previous independent studies. Nevertheless, two extracted scalars disagreed with these results. Furthermore, sensitivity analysis found that a relatively small coordinate system rotation of 5.5 degrees reversed the mediolateral null hypothesis rejection decision. Considering that the foot may adopt arbitrary postures in the horizontal plane, these sensitivity results suggest that non-negligible uncertainty may exist in mediolateral COP effects. As compared with COP scalar extraction, two key advantages of the vector-field approach are: (i) coordinate system independence, (ii) continuous statistical data reflecting the temporal extents of COP trajectory changes., Article, GAIT & POSTURE. 40(1):255-258 (2014)

The gait cycle is continuous, but for practical reasons one is often forced to analyze one or only a few adjacent cycles, for example in non-treadmill laboratory investigations and in fossilized footprint analysis. The nature of variability in long-term gait cycle dynamics has been well-investigated, but short-term variability, and specifically correlation, which are highly relevant to short gait bouts, have not. We presently tested for step-to-step autocorrelation in a total of 5243 plantar pressure (PP) distributions from ten subjects who walked at 1.1 m/s on an instrumented treadmill. Following spatial foot alignment, data were analyzed both from three points of interest (POI): heel, central metatarsals, and hallux, and for the foot surface as a whole, in a mass-univariate manner. POI results revealed low average step-to-step autocorrelation coefficients (r = 0.327 +/- 0.094; mean +/- st. dev.). Formal statistical testing of the whole-foot r distributions reached significance over an average of only 0.42 +/- 0.52% of the foot's surface, even for a highly conservative uncorrected threshold of p < 0.05. The common assumption, that short gait bouts consist of independent cycles, is therefore not refuted by the present PP results., Article, GAIT & POSTURE. 38(4):1054-1057 (2013)

We report a Holocene human and animal footprint site from the Namib Sand Sea, south of Walvis Bay, Namibia. Using these data, we explore intratrail footprint variability associated with small variations in substrate properties using a "whole foot" analytical technique developed for the studies in human ichnology. We demonstrate high levels of intratrail variability as a result of variations in grain size, depositional moisture content, and the degree of sediment disturbance, all of which determine the bearing capacity of the substrate. The two principal trails were examined, which had consistent stride and step lengths, and as such variations in print typology were primarily controlled by substrate rather than locomotor mechanics. Footprint typology varies with bearing capacity such that firm substrates show limited impressions associated with areas of peak plantar pressure, whereas softer substrates are associated with deep prints with narrow heels and reduced medial longitudinal arches. Substrates of medium bearing capacity give displacement rims and proximal movement of sediment, which obscures the true form of the medial longitudinal arch. A simple conceptual model is offered which summarizes these conclusions and is presented as a basis for further investigation into the control of substrate on footprint typology. The method, model, and results presented here are essential in the interpretation of any sites of greater paleoanthropological significance, such as recently reported from Ileret (1.5 Ma, Kenya; Bennett et al.: Science 323 (2009) 1197-1201).

During walking, variability in step parameters allows the body to adapt to changes in substrate or unexpected perturbations that may occur as the feet interface with the environment. Despite a rich literature describing biomechanical variability in step parameters, there are as yet no studies that consider variability at the body–environment interface. Here, we used pedobarographic statistical parametric mapping (pSPM) and two standard measures of variability, mean square error (m.s.e.) and the coefficient of variation (CV), to assess the magnitude and spatial variability in plantar pressure across a range of controlled walking speeds. Results by reduced major axis, and pSPM regression, revealed no consistent linear relationship between m.s.e. and speed or m.s.e. and Froude number. A positive linear relationship, however, was found between CV and walking speed and CV and Froude number. The spatial distribution of variability was highly disparate when assessed by m.s.e. and CV: relatively high variability was consistently confined to the medial and lateral forefoot when measured by m.s.e., while the forefoot and heel show high variability when measured by CV. In absolute terms, variability by CV was universally low (less than 2.5%). From these results, we determined that variability as assessed by m.s.e. is independent of speed, but dependent on speed when assessed by CV.

An animal's size is central to its ecology, yet remarkably little is known about the selective pressures that drive this trait. A particularly compelling example is how ancestral apes evolved large body mass in such a physically and energetically challenging environment as the forest canopy, where weight-bearing branches and lianas are flexible, irregular and discontinuous, and the majority of preferred foods are situated on the most flexible branches at the periphery of tree crowns. To date the issue has been intractable due to a lack of relevant fossil material, the limited capacity of the fossil record to reconstruct an animal's behavioural ecology and the inability to measure energy consumption in freely moving apes. We studied the oxygen consumption of parkour athletes while they traversed an arboreal-like course as an elite model ape, to test the ecomorphological and behavioural mechanisms by which a large-bodied ape could optimize its energetic performance during tree-based locomotion. Our results show that familiarity with the arboreal-like course allowed the athletes to substantially reduce their energy expenditure. Furthermore, athletes with larger arm spans and shorter legs were particularly adept at finding energetic savings. Our results flesh out the scanty fossil record to offer evidence that long, strong arms, broad chests and a strong axial system, combined with the frequent use of uniform branch-to-branch arboreal pathways, were critical to off-setting the mechanical and energetic demands of large mass in ancestral apes.

Living tree branches are almost impossible to snap. Some show “greenstick fracture”, breaking halfway across before splitting along their length, while others simply buckle. In this study we investigated the bending failure of coppice branches of three temperate angiosperm trees: ash, Fraxinus excelsior; hazel, Corylus avellana; and white willow, Salix alba. We carried out bending tests, and made a series of observations on the structure, density and tensile and compressive strength of their wood to understand the pattern of failure. The three species showed contrasting behaviour; willow buckled whereas ash showed clean greenstick fracture and hazel a more diffuse greenstick fracture. These differences could be related to their wood properties. Willow buckled because its light wood had very low transverse compressive strength, particularly tangentially and was crushed by transverse stresses. Though the other species yielded in longitudinal compression on the concave side, they ultimately failed in tension on the convex side when bent because their higher density wood resisted transverse compression better. However, the crack was diverted down the midline because of the low tangential tensile strength of their wood. Differences in fracture between ash and hazel are related to fine-scale differences in their wood anatomy and mechanics.

The maximum capability of a muscle can be estimated from simple measurements of muscle architecture such as muscle belly mass, fascicle length and physiological cross-sectional area. While the hindlimb anatomy of the non-human apes has been studied in some detail, a comparative study of the forelimb architecture across a number of species has never been undertaken. Here we present data from chimpanzees, bonobos, gorillas and an orangutan to ascertain if, and where, there are functional differences relating to their different locomotor repertoires and habitat usage. We employed a combination of analyses including allometric scaling and ANCOVAs to explore the data, as the sample size was relatively small and heterogeneous (specimens of different sizes, ages and sex). Overall, subject to possible unidentified, confounding factors such as age effects, it appears that the non-human great apes in this sample (the largest assembled to date) do not vary greatly across different muscle architecture parameters, even though they perform different locomotor behaviours at different frequencies. Therefore, it currently appears that the time spent performing a particular behaviour does not necessarily impose a dominating selective influence on the soft-tissue portion of the musculoskeletal system; rather, the overall consistency of muscle architectural properties both between and within the Asian and African apes strengthens the case for the hypothesis of a possible ancient shared evolutionary origin for orthogrady under compressive and ⁄ or suspensory loading in the great apes.

The distance that animals leap depends on their take-off angle and velocity. The velocity is generated solely by mechanical work during the push-off phase of standing-start leaps. Gibbons are capable of exceptional leaping performance, crossing gaps in the forest canopy exceeding 10 m, yet possess none of the adaptations possessed by specialist leapers synonymous with maximizing mechanical work. To understand this impressive performance, we recorded leaps of the gibbons exceeding 3.7 m. Gibbons perform more mass-specific work (35.4 J kg−1) than reported for any other species to date, accelerating to 8.3 ms−1in a single movement and redefining our estimates of work performance by animals. This energy (enough for a 3.5 m vertical leap) is 60 per cent higher than that achieved by galagos, which are renowned for their remarkable leaping performance. The gibbons' unusual morphology facilitates a division of labour among the hind limbs, forelimbs and trunk, resulting in modest power requirements compared with more specialized leapers.

By relating an animal’s morphology to its functional role and the behaviours performed, we can further develop our understanding of the selective factors and constraints acting on the adaptations of great apes. Comparison of muscle architecture between different ape species, however, is difficult because only small sample sizes are ever available. Further, such samples are often comprised of different age–sex classes, so studies have to rely on scaling techniques to remove body mass differences. However, the reliability of such scaling techniques has been questioned. As datasets increase in size, more reliable statistical analysis may eventually become possible. Here we employ geometric and allometric scaling techniques, and ancovas (a form of general linear model, GLM) to highlight and explore the different methods available for comparing functional morphology in the non-human great apes. Our results underline the importance of regressing data against a suitable body size variable to ascertain the relationship (geometric or allometric) and of choosing appropriate exponents by which to scale data. ancova models, while likely to be more robust than scaling for species comparisons when sample sizes are high, suffer from reduced power when sample sizes are low. Therefore, until sample sizes are radically increased it is preferable to include scaling analyses along with ancovas in data exploration. Overall, the results obtained from the different methods show little significant variation, whether in muscle belly mass, fascicle length or physiological cross-sectional area between the different species. This may reflect relatively close evolutionary relationships of the non-human great apes; a universal influence on morphology of generalised orthograde locomotor behaviours or, quite likely, both.

In an arboreal habitat, primates have to cope with a complex meshwork of flexible supports in order to obtain food, find mates and avoid predators. To understand how animals interact with such complex environments we can study their positional behaviour. However, due to the intricate variation in locomotion and posture it can be difficult to capture details such as limb use (i.e. weight and balance), limb flexion and substrate use. This paper presents a suitable method replicable for any primate species, based on the movement notation technique, Sutton Movement Writing (SMW), aiming to record the spatial arrangement of limbs during positional behaviours on multiple, compliant supports. This method was piloted during a year-long field study of wild orangutans (Pongo abelii) and validated and tested for inter- and intraobserver reliability using videos from the field. Overall, SMW shows considerable promise for increasing the resolution with which positional behaviours can be recorded under field conditions and provides a way to extract numerical data for use in statistical analyses. This will facilitate our understanding of how behaviours vary in response to the environment, and the capabilities of primates to perform key tasks in their distinct niches.

The full publication ofArdipithecus ramidushas particular importance for the origins of hominin bipedality, and strengthens the growing case for an arboreal origin. Palaeontological techniques however inevitably concentrate on details of fragmentary postcranial bones and can benefit from a whole-animal perspective. This can be provided by field studies of locomotor behaviour, which provide a real-world perspective of adaptive context, against which conclusions drawn from palaeontology and comparative osteology may be assessed and honed. Increasingly sophisticated dynamic modelling techniques, validated against experimental data for living animals, offer a different perspective where evolutionary and virtual ablation experiments, impossible for living mammals, may be runin silico, and these can analyse not only the interactions and behaviour of rigid segments but increasingly the effects of compliance, which are of crucial importance in guiding the evolution of an arboreally derived lineage.

The plantar aponeurosis (PA), in spanning the whole length of the plantar aspect of the foot, is clearly identified as one of the key structures that is likely to affect compliance and stability of the longitudinal arch. A recent study performed in our laboratory showed that tension/elongation in the PA can be predicted from the kinematics of the segments to which the PA is attached. In the present investigation, stereophotogrammetry and inverse kinematics were employed to shed light on the mechanics of the longitudinal arch and its main passive stabilizer, the PA, in relation to walking speed. When compared with a neutral unloaded position, the medial longitudinal arch underwent greater collapse during the weight-acceptance phase of stance at higher walking speed (0.1°±1.9° in slow walking; 0.9°±2.6° in fast walking; P = 0.0368). During late stance the arch was higher (3.4°±3.1° in slow walking; 2.8°±2.7° in fast walking; P = 0.0227) and the metatarsophalangeal joints more dorsiflexed (e.g. at the first metatarsophalangeal joint, 52°±5° in slow walking; 64°±4° in fast walking; P < 0.001) during fast walking. Early-stance tension in the PA increased with speed, whereas maximum tension during late stance did not seem to be significantly affected by walking speed. Although, on the one hand, these results give evidence for the existence of a pre-heel-strike, speed-dependent, arch-stiffening mechanism, on the other hand they suggest that augmentation of arch height in late stance is enhanced by higher forces exerted by the intrinsic muscles on the plantar aspect of the foot when walking at faster speeds.

Evidence has frequently been reported of modifications in gait patterns within the lower limb related to the cadence of walking. Most reports have concerned relationships between cadence and kinematic and the kinetic changes occurring in the main joints and muscles of the lower limb as a whole. The aim of the present study was to assess whether significant changes are also measurable in kinematics of the foot segments. An existing 15 marker-set protocol allowed a four-segment foot and shank model to be defined for relative rotations between the segments to be calculated. Stereophotogrammetry was employed to record marker position data from ten subjects walking at three cadences. The slow- and normal cadence datasets showed similar profiles of joint rotation in three anatomical planes, but significant differences were found between these and the fast cadence. At all joints, frame-by-frame statistical analysis revealed increased dorsiflexion from heel-strike to midstance (p < 0.05) and increased plantarflexion from midstance to toe-off (p < 0.05) with increasing cadence. From foot-flat to heel-rise, the fast cadence kinematic data showed a decreased range of motion in the sagittal-plane between forefoot and rearfoot (3.2 degrees +/- 1.2 degrees at slow cadence; 2.0 degrees +/- 0.8 degrees at fast cadence; p < 0.05). The cadences imposed and the multisegment protocol revealed significant kinematic changes in the joints of the foot during barefoot walking.

Napier and Walker’s (1967) locomotor category of vertical clinging and leaping (VC&L) is one of the most familiar in primatology, and tarsiers are probably the most morphologically specialized of its membership. However, the link between vertical clinging and leaping remains unelucidated. We attempt to do so by reanalysis of Crompton’s 1985 and 1986 field observations of locomotion and habitat use in Tarsius bancanus, using loglinear modeling. Loglinear modeling is better suited to the categorical variables used in many field studies than more traditional statistics, such as ANOVA, developed for continuous variables. We show that climbing, as well as leaping, is one of the predominant forms of locomotion, and that all other things being equal, tarsiers tend to take off from, and land on, similar sized supports, which suggests that the following findings are not likely to be a result of substrate availability alone. Small body size lead to a prediction that tarsiers should leap down but climb up: this was not sustained: rather leaps tend to be level, and climbing accounts for more height loss than randomly expected. However, a prediction that to avoid energy loss to the substrate, the tarsiers should show a preference for large diameter supports for takeoff when leaping longer distances was supported, although tarsiers do not avoid moderately compliant supports. The prediction from ballistic principles that the longest leaps should start from high-angled supports was only weakly sustained, but low-angled supports tend to be strongly associated with short leaps, suggesting that such supports do not facilitate 45° takeoff trajectories. However, tarsiers displayed a preference for landing on medium-sized supports when leaping long distances, suggesting a preference for balancing the need for stability with minimizing musculoskeletal shock.

Gibbons are skilled brachiators but they are also highly capable leapers, crossing distances in excess of 10 m in the wild. Despite this impressive performance capability, no detailed biomechanical studies of leaping in gibbons have been undertaken to date. We measured ground reaction forces and derived kinematic parameters from high-speed videos during gibbon leaps in a captive zoo environment. We identified four distinct leap types defined by the number of feet used during take-off and the orientation of the trunk, orthograde single-footed, orthograde two-footed, orthograde squat, and pronograde single-footed leaps. The center of mass trajectories of three of the four leap types were broadly similar, with the pronograde single-footed leaps exhibiting less vertical displacement of the center of mass than the other three types. Mechanical energy at take-off was similar in all four leap types. The ratio of kinetic energy to mechanical energy was highest in pronograde single-footed leaps and similar in the other three leap types. The highest mechanical work and power were generated during orthograde squat leaps. Take-off angle decreased with take-off velocity and the hind limbs showed a proximal to distal extension sequence during take-off. In the forelimbs, the shoulder joints were always flexed at take-off, while the kinematics of the distal joints (elbow and wrist joints) were variable between leaps. It is possible that gibbons may utilize more metabolically expensive orthograde squat leaps when a safe landing is uncertain, while more rapid (less expensive) pronograde single-footed leaps might be used during bouts of rapid locomotion when a safe landing is more certain.

Computational steering is an investigative paradigm whereby the parameters of a running program can be altered according to what is seen in the currently visualized results of the simulation. For certain problems, interactive computation brings specific benefits: parameter sweeps can be completed more efficiently by quickly identifying combinations of input values that yield nonsensical results; 'what-if' studies may elucidate a number of related hypotheses without computing all of them from scratch; even simply tracking the development of a computation on-line may allow its early termination (with consequent resource savings) if it turns out its set-up was flawed for some reason. Having first come to prominence in the 1990s, the take-up of steering has accelerated in recent years: high performance computer (HPC) facilites have become more widely available, usage of the Grid is increasing, and visualization is experiencing take-up beyond the walls of the pioneering graphics laboratories that first nurtured it. All these factors combine to expand the range and difficulty of the scientific problems that can be tackled by steering. However, this benefit comes at the cost of increasing hardware and software complexity which itself may defeat further progress. This paper discusses these issues and, in the light of some users' recent experiences, charts the challenges that still face us.

Gibbons utilize a number of locomotor modes in the wild, including bipedalism, leaping and, most of all, brachiation. Each locomotor mode puts specific constraints on the morphology of the animal; in some cases these may be complementary, whereas in others they may conflict. Despite several studies of the locomotor biomechanics of gibbons, very little is known about the musculoskeletal architecture of the limbs. In this study, we present quantitative anatomical data of the hind limb for four species of gibbon (Hylobates lar, H. moloch, H. pileatus and Symphalangus syndactylus). Muscle mass and fascicle lengths were obtained from all of the major hind limb muscles and the physiological cross-sectional area was calculated and scaled to remove the effect of body size. The results clearly indicate that, for all of the species studied, the major hip, knee and ankle extensors are short-fascicled and pennate. The major hip and knee flexors, however, are long-fascicled, parallel muscles with relatively small physiological cross-sectional areas. We hypothesize that the short-fascicled muscles could be coupled with a power-amplifying mechanism and are predominantly useful in leaping. The long-fascicled knee and hip flexors are adapted for a wide range of joint postures and can play a role in flexing the legs during brachiation. ispartof: Journal of anatomy vol:215 issue:4 pages:383-400 ispartof: location:England status: published

SUMMARY In the present study we have estimated the temporal elongation of the plantar aponeurosis (PA) during normal walking using a subject-specific multi-segment rigid-body model of the foot. As previous studies have suggested that muscular forces at the ankle can pre-load the PA prior to heel-strike,the main purpose of the current study was to test, through modelling, whether there is any tension present in the PA during early stance phase. Reflective markers were attached to bony landmarks to track the kinematics of the calcaneus, metatarsus and toes during barefoot walking. Ultrasonography measurements were performed on three subjects to determine both the location of the origin of the PA on the plantar aspect of the calcaneus, and the radii of the metatarsal heads. Starting with the foot in a neutral, unloaded position, inverse kinematics allowed calculation of the tension in the five slips of the PA during the whole duration of the stance phase. The results show that the PA experienced tension significantly above rest during early stance phase in all subjects (P

Image registration, the process of transforming images such that homologous structures optimally overlap, provides the pre-processing foundation for pixel-level functional image analysis. The purpose of this study was to compare the performances of seven methods of within-subjects pedobarographic image registration: (1) manual, (2) principal axes, (3) centre of pressure trajectory, (4) mean squared error, (5) probability-weighted variance, (6) mutual information, and (7) exclusive OR. We assumed that foot-contact geometry changes were negligibly small trial-to-trial and thus that a rigid-body transformation could yield optimum registration performance. Thirty image pairs were randomly selected from our laboratory database and were registered using each method. To compensate for inter-rater variability, the mean registration parameters across 10 raters were taken as representative of manual registration. Registration performance was assessed using four dissimilarity metrics (#4-7 above). One-way MANOVA found significant differences between the methods (p

Based on our knowledge of locomotor biomechanics and ecology we predict the locomotion and posture of the last common ancestors of (a) great and lesser apes and their close fossil relatives (hominoids); (b) chimpanzees, bonobos and modern humans (hominines); and (c) modern humans and their fossil relatives (hominins). We evaluate our propositions against the fossil record in the context of a broader review of evolution of the locomotor system from the earliest hominoids of modern aspect (crown hominoids) to early modern Homo sapiens. While some early East African stem hominoids were pronograde, it appears that the adaptations which best characterize the crown hominoids are orthogrady and an ability to abduct the arm above the shoulder - rather than, as is often thought, manual suspension sensu stricto. At 7-9 Ma (not much earlier than the likely 4-8 Ma divergence date for panins and hominins, see Bradley, 2008) there were crown hominoids in southern Europe which were adapted to moving in an orthograde posture, supported primarily on the hindlimb, in an arboreal, and possibly for Oreopithecus, a terrestrial context. By 7 Ma, Sahelanthropus provides evidence of a Central African hominin, panin or possibly gorilline adapted to orthogrady, and both orthogrady and habitually highly extended postures of the hip are evident in the arboreal East African protohominin Orrorin at 6 Ma. If the traditional idea that hominins passed through a terrestrial 'knuckle-walking' phase is correct, not only does it have to be explained how a quadrupedal gait typified by flexed postures of the hindlimb could have preadapted the body for the hominin acquisition of straight-legged erect bipedality, but we would have to accept a transition from stem-hominoid pronogrady to crown hominoid orthogrady, back again to pronogrady in the African apes and then back to orthogrady in hominins. Hand-assisted arboreal bipedality, which is part of a continuum of orthograde behaviours, is used by modern orangutans to forage among the small branches at the periphery of trees where the core hominoid dietary resource, ripe fruit, is most often to be found. Derivation of habitual terrestrial bipedality from arboreal hand-assisted bipedality requires fewer transitions, and is also kinematically and kinetically more parsimonious. ispartof: Journal of anatomy vol:212 issue:4 pages:501-543 ispartof: location:ENGLAND, St Annes Coll, Oxford status: published

This study investigates the relation between walking speed and the distribution of peak plantar pressure and compares a traditional ten-region subsampling (10RS) technique with a new technique: pedobarographic statistical parametric mapping (pSPM). Adapted from cerebral fMRI methodology, pSPM is a digital image processing technique that registers foot pressure images such that homologous structures optimally overlap, thereby enabling statistical tests to be conducted at the pixel level. Following previous experimental protocols, we collected pedobarographic records from 10 subjects walking at three different speeds: slow, normal, and fast. Walking speed was recorded and correlated with the peak pressures extracted from the 10 regions, and subsequently with the peak pixel data extracted after pSPM preprocessing. Both methods revealed significant positive correlation between peak plantar pressure and walking speed over the rearfoot and distal forefoot after Bonferroni correction for multiple comparisons. The 10RS analysis found positive correlation in the midfoot and medial proximal forefoot, but the pixel data exhibited significant negative correlation throughout these regions (p5x10(-5)). Comparing the statistical maps from the two approaches shows that subsampling may conflate pressure differences evident in pixel-level data, obscuring or even reversing statistical trends. The negative correlation observed in the midfoot implies reduced longitudinal arch collapse with higher walking speeds. We infer that this results from pre- or early-stance phase muscle activity and speculate that preferred walking speed reflects, in part, a balance between the energy required to tighten the longitudinal arch and the apparent propulsive benefits of the stiffened arch.

Human bipedalism is commonly thought to have evolved from a quadrupedal terrestrial precursor, yet some recent paleontological evidence suggests that adaptations for bipedalism arose in an arboreal context. However, the adaptive benefit of arboreal bipedalism has been unknown. Here we show that it allows the most arboreal great ape, the orangutan, to access supports too flexible to be negotiated otherwise. Orangutans react to branch flexibility like humans running on springy tracks, by increasing knee and hip extension, whereas all other primatesdothe reverse. Human bipedalism is thus less an innovation than an exploitation of a locomotor behavior retained from the common great ape ancestor.

Within the forest canopy, the shortest gaps between tree crowns lie between slender terminal branches. While the compliance of these supports has previously been shown to increase the energetic cost of gap crossing in arboreal animals (e.g. Alexander 1991Z. Morphol. Anthropol.78, 315–320; Demeset al. 1995Am. J. Phys. Anthropol.96, 419–429), field observations suggest that some primates may be able to use support compliance to increase the energetic efficiency of locomotion. Here, we calculate the energetic cost of alternative methods of gap crossing in orangutans (Pongo abelii). Tree sway (in which orangutans oscillate a compliant tree trunk with increasing magnitude to bridge a gap) was found to be less than half as costly as jumping, and an order of magnitude less costly than descending the tree, walking to the vine and climbing it. Observations of wild orangutans suggest that they actually use support compliance in many aspects of their locomotor behaviour. This study seems to be the first to show that elastic compliance in arboreal supports can be used to reduce the energetic cost of gap crossing.