The common chimpanzee (Pan troglodytes) is a facultative biped and our closest living relative. As such, the musculoskeletal anatomies of their pelvis and hind limbs have long provided a comparative context for studies of human and fossil hominin locomotion. Yet, how the chimpanzee pelvis and hind limb actually move during bipedal walking is still not well defined. Here, we describe the three-dimensional (3-D) kinematics of the pelvis, hip, knee and ankle during bipedal walking and compare those values to humans walking at the same dimensionless and dimensional velocities. The stride-to-stride and intraspecific variations in 3-D kinematics were calculated using the adjusted coefficient of multiple correlation. Our results indicate that humans walk with a more stable pelvis than chimpanzees, especially in tilt and rotation. Both species exhibit similar magnitudes of pelvis list, but with segment motion that is opposite in phasing. In the hind limb, chimpanzees walk with a more flexed and abducted limb posture, and substantially exceed humans in the magnitude of hip rotation during a stride. The average stride-to-stride variation in joint and segment motion was greater in chimpanzees than humans, while the intraspecific variation was similar on average. These results demonstrate substantial differences between human and chimpanzee bipedal walking, in both the sagittal and non-sagittal planes. These new 3-D kinematic data are fundamental to a comprehensive understanding of the mechanics, energetics and control of chimpanzee bipedalism.

The longitudinal arch of the human foot is commonly thought to reduce midfoot joint motion to convert the foot into a rigid lever during push off in bipedal walking. In contrast, African apes have been observed to exhibit midfoot dorsiflexion following heel lift during terrestrial locomotion, presumably due to their possession of highly mobile midfoot joints. This assumed dichotomy between human and African ape midfoot mobility has recently been questioned based on indirect assessments of in vivo midfoot motion, such as plantar pressure and cadaver studies; however, direct quantitative analyses of African ape midfoot kinematics during locomotion remain scarce. Here, we used high-speed motion capture to measure three-dimensional foot kinematics in two male chimpanzees and five male humans walking bipedally at similar dimensionless speeds. We analyzed 10 steps per chimpanzee subject and five steps per human subject, and compared ranges of midfoot motion between species over stance phase, as well as within double- and single-limb support periods. Contrary to expectations, humans used a greater average range of midfoot motion than chimpanzees over the full duration of stance. This difference was driven by humans' dramatic plantarflexion and adduction of the midfoot joints during the second double-limb support period, which likely helps the foot generate power during push off. However, chimpanzees did use slightly but significantly more midfoot dorsiflexion than humans in the single limb-support period, during which heel lift begins. These results indicate that both stiffness and mobility are important to longitudinal arch function, and that the human foot evolved to utilize both during push off in bipedal walking. Thus, the presence of human-like midfoot joint morphology in fossil hominins should not be taken as indicating foot rigidity, but may signify the evolution of pedal anatomy conferring enhanced push off mechanics.

Humans diverged from apes (chimpanzees, specifically) toward the end of the Miocene ~9.3 million to 6.5 million years ago. Understanding the origins of the human lineage (hominins) requires reconstructing the morphology, behavior, and environment of the chimpanzee-human last common ancestor. Modern hominoids (that is, humans and apes) share multiple features (for example, an orthograde body plan facilitating upright positional behaviors). However, the fossil record indicates that living hominoids constitute narrow representatives of an ancient radiation of more widely distributed, diverse species, none of which exhibit the entire suite of locomotor adaptations present in the extant relatives. Hence, some modern ape similarities might have evolved in parallel in response to similar selection pressures. Current evidence suggests that hominins originated in Africa from Miocene ape ancestors unlike any living species.

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Large-scale interspecific studies of mammals ranging between 0.04–280 kg have shown that larger animals walk with more extended limb joints. Within a taxon or clade, however, the relationship between body size and joint posture is less straightforward. Factors that may affect the lack of congruence between broad and narrow phylogenetic analyses of limb kinematics include limited sampling of (1) ranges of body size, and/or (2) numbers of individuals. Unfortunately, both issues are inherent in laboratory-based or zoo locomotion research. In this study, we examined the relationship between body mass and elbow and knee joint angles (our proxies of fore- and hind limb posture, respectively) in a cross-sectional ontogenetic sample of wild chacma baboons (Papio hamadryas ursinus) habituated in the De Hoop Nature Reserve, South Africa. Videos were obtained from 33 individuals of known age (12 to ≥108 months) and body mass (2–29.5 kg) during walking trials. Results show that older, heavier baboons walk with significantly more extended knee joints but not elbow joints. This pattern is consistent when examining only males, but not within the female sample. Heavier, older baboons also display significantly less variation in their hind limb posture compared to lighter, young animals. Thus, within this ontogenetic sample of a single primate species spanning an order of magnitude in body mass, hind limb posture exhibited a postural scaling phenomenon while the forelimbs did not. These findings may further help explain 1) why younger mammals (including baboons) tend to have relatively stronger bones than adults, and 2) why humeri appear relatively weaker than femora (in at least baboons). Finally, this study demonstrates how field-acquired kinematics can help answer fundamental biomechanical questions usually addressed only in animal gait laboratories.

The origin and evolution of knuckle-walking has long been a key focus in understanding African ape, including human, origins. Yet, despite numerous studies documenting morphological characteristics potentially associated with knuckle-walking, little quantitative three-dimensional (3-D) data exist of forelimb motion during knuckle-walking. Nor do any comparative 3-D data exist for hand postures used during quadrupedalism in monkeys. This lack of data has limited the testability of proposed adaptations for knuckle-walking in African apes. This study presents the first 3-D kinematic data of the wrist, hand and metacarpophalangeal joints during knuckle-walking in chimpanzees and in macaques using digitigrade and palmigrade hand postures. These results clarify the unique characteristics of, and commonalities between, knuckle-walking and digitigrady/palmigrady in multiple planes of motion. Notably, chimpanzees utilized more wrist ulnar deviation than any macaque hand posture. Maximum extension of the chimpanzee wrist was slight (5–20 deg) and generally overlapped with macaque digitigrady. Metacarpophalangeal joint motion displayed distinct differences between digits in both species, likely related to the timing of force application. These data also reveal that maximum metacarpophalangeal extension angles during knuckle-walking (26–59 deg) were generally higher than previously considered. In macaques, maximum metacarpophalangeal extension during digitigrady and palmigrady overlapped for most digits, highlighting additional complexity in the interpretation of skeletal features that may be related to limiting metacarpophalangeal motion. Most importantly, however, these new 3-D data serve as a fundamental dataset with which evaluation of proposed musculoskeletal adaptations for knuckle-walking can be tested.

Humans are unique among apes and other primates in the musculoskeletal design of their lower back and pelvis. While the last common ancestor of the lineages has long been thought to be 'African ape-like', including in its lower back and ilia design, recent descriptions of early hominin and Miocene ape fossils have led to the proposal that its lower back and ilia were more similar to those of some Old World monkeys, such as macaques. Here, we compared three-dimensional kinematics of the pelvis and hind/lower limbs of bipedal macaques, chimpanzees and humans walking at similar dimensionless speeds to test the effects of lower back and ilia design on gait. Our results indicate that locomotor kinematics of bipedal macaques and chimpanzees are remarkably similar, with both species exhibiting greater pelvis motion and more flexed, abducted hind limbs than humans during walking. Some differences between macaques and chimpanzees in pelvis tilt and hip abduction were noted, but they were small in magnitude; larger differences were observed in ankle flexion. Our results suggest that if and diverged from a common ancestor whose lower back and ilia were either 'African ape-like' or more 'Old World monkey-like', at its origin, the hominin walking stride likely involved distinct (i.e. non-human-like) pelvis motion on flexed, abducted hind limbs.

During terrestrial locomotion, chimpanzees exhibit dorsiflexion of the midfoot between midstance and toe-off of stance phase, a phenomenon that has been called the "midtarsal break." This motion is generally absent during human bipedalism, and in chimpanzees is associated with more mobile foot joints than in humans. However, the contribution of individual foot joints to overall foot mobility in chimpanzees is poorly understood, particularly on the medial side of the foot. The talonavicular (TN) and calcaneocuboid (CC) joints have both been suggested to contribute significantly to midfoot mobility and to the midtarsal break in chimpanzees. To evaluate the relative magnitude of motion that can occur at these joints, we tracked skeletal motion of the hindfoot and midfoot during passive plantarflexion and dorsiflexion manipulations using cineradiography. The sagittal plane range of motion was 38 ± 10° at the TN joint and 14 ± 8° at the CC joint. This finding indicates that the TN joint is more mobile than the CC joint during ankle plantarflexion-dorsiflexion. We suggest that the larger range of motion at the TN joint during dorsiflexion is associated with a rotation (inversion-eversion) across the transverse tarsal joint, which may occur in addition to sagittal plane motion.