The long limb bones of the StW 573 Australopithecus skeleton from Sterkfontein Member 2: Descriptions and proportions.

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 chimp/human last common ancestor (CLCA); (2) indication of some degree of climbing ability; (3) allometry. Recent discoveries of other partial skeletons of Australopithecus, such as those of Australopithecus sediba (MH1 and MH2) and Australopithecus 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) skeleton 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. 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 a purely allometric explanation for A.L. 288-1 limb proportions. In fact, our multivariate allometric analysis suggests that limb lengths of Australopithecus, as represented by StW 573 and A.L. 288-1, exhibit a significantly different (p < 0.001) allometric pattern than that which typifies modern humans and African apes. Like some previous analyses, our results also suggest that hominin limb evolution occurred in two stages with: first, a modest increase in lower limb length and a concurrent shortening of the antebrachium between Ardipithecus and Australopithecus, followed by a considerable lengthening of the lower limb along with a decrease of both upper limb elements occurring between Australopithecus and Homo sapiens.

A Three-Dimensional Musculoskeletal Model of the Western Lowland Gorilla Foot: Examining Muscle Torques and Function.

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.

A 3D musculoskeletal model of the western lowland gorilla hind limb: moment arms and torque of the hip, knee and ankle.

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.

Practice makes perfect: Performance optimisation in 'arboreal' parkour athletes illuminates the evolutionary ecology of great ape anatomy.

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.

Locomotor flexibility in Lepilemur explained by habitat and biomechanics.

There are at present few comparable studies of lemur locomotion in the wild. This has unfortunately 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 Ankarafantsika 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 orientation, heights, and distances traveled were all collected. 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 supports ≤5 cm in diameter and L. edwardsi >5 cm in diameter.

Nest-building orangutans demonstrate engineering know-how to produce safe, comfortable beds.

Nest-building orangutans must daily build safe and comfortable nest structures in the forest canopy and do this quickly and effectively using the branches that surround them. This study aimed to investigate the mechanical design and architecture of orangutan nests and determine the degree of technical sophistication used in their construction. We measured the whole nest compliance and the thickness of the branches used and recorded the ways in which the branches were fractured. Branch samples were also collected from the nests and subjected to three-point bending tests to determine their mechanical properties. We demonstrated that the center of the nest is more compliant than the edges; this may add extra comfort and safety to the structure. During construction orangutans use the fact that branches only break half-way across in "greenstick" fracture to weave the main nest structure. They choose thicker branches with greater rigidity and strength to build the main structure in this way. They then detach thinner branches by following greenstick fracture with a twisting action to make the lining. These results suggest that orangutans exhibit a degree of technical knowledge and choice in the construction of nests.

Locomotor Ecology of Propithecus verreauxi in Kirindy Mitea National Park.

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.

Analysis of joint force and torque for the human and non-human ape foot during bipedal walking with implications for the evolution of the foot.

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.

The evolution of compliance in the human lateral mid-foot.

Fossil evidence for longitudinal arches in the foot is frequently used to constrain the origins of terrestrial bipedality in human ancestors. This approach rests on the prevailing concept that human feet are unique in functioning with a relatively stiff lateral mid-foot, lacking the significant flexion and high plantar pressures present in non-human apes. This paradigm has stood for more than 70 years but has yet to be tested objectively with quantitative data. Herein, we show that plantar pressure records with elevated lateral mid-foot pressures occur frequently in healthy, habitually shod humans, with magnitudes in some individuals approaching absolute maxima across the foot. Furthermore, the same astonishing pressure range is present in bonobos and the orangutan (the most arboreal great ape), yielding overlap with human pressures. Thus, while the mean tendency of habitual mechanics of the mid-foot in healthy humans is indeed consistent with the traditional concept of the lateral mid-foot as a relatively rigid or stabilized structure, it is clear that lateral arch stabilization in humans is not obligate and is often transient. These findings suggest a level of detachment between foot stiffness during gait and osteological structure, hence fossilized bone morphology by itself may only provide a crude indication of mid-foot function in extinct hominins. Evidence for thick plantar tissues in Ardipithecus ramidus suggests that a human-like combination of active and passive modulation of foot compliance by soft tissues extends back into an arboreal context, supporting an arboreal origin of hominin bipedalism in compressive orthogrady. We propose that the musculoskeletal conformation of the modern human mid-foot evolved under selection for a functionally tuneable, rather than obligatory stiff structure.

Holocene footprints in Namibia: the influence of substrate on footprint variability.

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).

Functional adaptations in the forelimb muscles of non-human great apes.

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 extraordinary athletic performance of leaping gibbons.

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(-1) in 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.

Orangutans employ unique strategies to control branch flexibility.

Orangutans are the largest habitually arboreal mammal. For them, as for all arboreal mammals, access to the abundant fruits and narrowest gaps found among the thin peripheral branches of tree crowns poses considerable safety risks and energetic demands. Most arboreal primates use flexed-limb postures to minimize problems caused by branch compliance and instability. Here, we show that Sumatran orangutans employ unique locomotor strategies to control compliance and allow access to the terminal branch niche for feeding and gap crossing. We calculated a "stiffness score," which is a measure of the flexibility of the supports on which orangutans moved. We found that certain locomotor behaviors clearly are associated with the most compliant supports; these behaviors appear to lack regular limb sequences, which serves to avoid the risk of resonance in branch sway caused by high-frequency, patterned gait. Balance and increased stability are achieved through long contact times between multiple limbs and supports and a combination of pronograde (horizontal) and orthograde (vertical) body postures, used both above branches and in suspension underneath them. Overall, adult females seem to be the most conservative in their travel, selecting more solid and secure supports than males and adolescents. These results have implications for understanding locomotor diversity in fossil and extant apes and for orangutan conservation and reintroduction programs.

Hindlimb muscle architecture in non-human great apes and a comparison of methods for analysing inter-species variation.

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.

The effect of substrate compliance on the biomechanics of gibbon leaps.

The storage and recovery of elastic strain energy in the musculoskeletal systems of locomoting animals has been extensively studied, yet the external environment represents a second potentially useful energy store that has often been neglected. Recent studies have highlighted the ability of orangutans to usefully recover energy from swaying trees to minimise the cost of gap crossing. Although mechanically similar mechanisms have been hypothesised for wild leaping primates, to date no such energy recovery mechanisms have been demonstrated biomechanically in leapers. We used a setup consisting of a forceplate and two high-speed video cameras to conduct a biomechanical analysis of captive gibbons leaping from stiff and compliant poles. We found that the gibbons minimised pole deflection by using different leaping strategies. Two leap types were used: slower orthograde leaps and more rapid pronograde leaps. The slower leaps used a wider hip joint excursion to negate the downward movement of the pole, using more impulse to power the leap, but with no increase in work done on the centre of mass. Greater hip excursion also minimised the effective leap distance during orthograde leaps. The more rapid leaps conversely applied peak force earlier in stance where the pole was effectively stiffer, minimising deflection and potential energy loss. Neither leap type appeared to usefully recover energy from the pole to increase leap performance, but the gibbons demonstrated an ability to best adapt their leap biomechanics to counter the negative effects of the compliant pole.

Muscle moment arms of the gibbon hind limb: implications for hylobatid locomotion.

Muscles facilitate skeletal movement via the production of a torque or moment about a joint. The magnitude of the moment produced depends on both the force of muscular contraction and the size of the moment arm used to rotate the joint. Hence, larger muscle moment arms generate larger joint torques and forces at the point of application. The moment arms of a number of gibbon hind limb muscles were measured on four cadaveric specimens (one Hylobates lar, one H. moloch and two H. syndactylus). The tendon travel technique was used, utilizing an electro-goniometer and a linear voltage displacement transducer. The data were analysed using a technique based on a differentiated cubic spline and normalized to remove the effect of body size. The data demonstrated a functional differentiation between voluminous muscles with short fascicles having small muscle moment arms and muscles with longer fascicles and comparatively smaller physiological cross-sectional area having longer muscle moment arms. The functional implications of these particular configurations were simulated using a simple geometric fascicle strain model that predicts that the rectus femoris and gastrocnemius muscles are more likely to act primarily at their distal joints (knee and ankle, respectively) because they have short fascicles. The data also show that the main hip and knee extensors maintain a very small moment arm throughout the range of joint angles seen in the locomotion of gibbons, which (coupled to voluminous, short-fascicled muscles) might help facilitate rapid joint rotation during powerful movements.

Keep your head down: Maintaining gait stability in challenging conditions.

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.

Look out: an exploratory study assessing how gaze (eye angle and head angle) and gait speed are influenced by surface complexity.

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.

Physical and perceptual measures of walking surface complexity strongly predict gait and gaze behaviour.

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.

The atlas of StW 573 and the late emergence of human-like head mobility and brain metabolism.

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.