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The initiation of a walking step with a heel strike is a defining characteristic of humans and great apes but is rarely found in other mammals. Despite the considerable importance of heel strike to an understanding of human locomotor evolution, no one has explicitly tested the fundamental mechanical question of why great apes use a heel strike. In this report, we test two hypotheses (1) that heel strike is a function of hip protraction and/or knee extension and (2) that short-legged apes with a midfoot that dorsiflexes at heel lift and long digits for whom digitigrady is not an option use heel-strike plantigrady. This strategy increases hip translation while potentially moderating the cost of redirecting the center of mass ('collisional costs') during stance via rollover along the full foot from the heel to toes. We quantified hind limb kinematics and relative hip translation in ten species of primates, including lemurs, terrestrial and arboreal monkeys, chimpanzees, and gorillas. Chimpanzees and gorillas walked with relatively extended knees but only with moderately protracted hips or hind limbs, partially rejecting the first hypothesis. Nonetheless, chimpanzees attained relative hip translations comparable with those of digitigrade primates. Heel-strike plantigrady may be a natural result of a need for increased hip translations when forelimbs are relatively long and digitigrady is morphologically restricted. In addition, foot rollover from the heel to toe in large, short-legged apes may reduce energetic costs of redirecting the center of mass at the step-to-step transition as it appears to do in humans. Heel strike appears to have been an important mechanism for increasing hip translation, and possibly reducing energetic costs, in early hominins and was fundamental to the evolution of the modern human foot and human bipedalism.

Arm-swinging is a locomotor mode observed only in primates, in which the hindlimbs no longer have a weight bearing function and the forelimbs must propel the body forward and support the entirety of the animal's mass. It has been suggested that the evolution of arm-swinging was preceded by a shift to inverted quadrupedal walking for purposes of feeding and balance, yet little is known about the mechanics of limb use during inverted quadrupedal walking. In this study, we test whether the mechanics of inverted quadrupedal walking make sense as precursors to arm-swinging and whether there are fundamental differences in inverted quadrupedal walking in primates compared to non-primate mammals that would explain the evolution of arm-swinging in primates only. Based on kinetic limb-loading data collected during inverted quadrupedal walking in primates (seven species) and non-primate mammals (three species), we observe that in primates the forelimb serves as the primary propulsive and weight bearing limb. Additionally, heavier individuals tend to support a greater distribution of body weight on their forelimbs than lighter ones. These kinetic patterns are not observed in non-primate mammals. Based on these findings, we propose that the ability to adopt arm-swinging is fairly simple for relatively large-bodied primates and merely requires the animal to release its grasping foot from the substrate. This study fills an important gap concerning the origins of arm-swinging and illuminates previously unknown patterns of primate locomotor evolution.

Facultative bipedalism during load transport in nonhuman primates has been argued to be an important behavior potentially leading to the evolution of obligate, extended limb bipedalism. Understanding the biomechanics of such behavior may lead to insights about associated morphology, which may translate to interpretation of features in the fossil record. Some populations of bearded capuchin monkeys (Sapajus libidinosus) spontaneously carry heavy loads bipedally during foraging activities. This study provides the first data on all three components of ground reaction force for spontaneous bipedalism during load carriage in a nonhuman primate. Five individual S. libidinosus (mean body mass = 2.4 kg +/- 0.96) were videorecorded during bipedalism while carrying a stone (0.93 kg) under natural conditions. A force plate was embedded in the path of the monkeys. Spatiotemporal and force data for all three components of the ground reaction force were recorded for 28 steps. Capuchins exhibited a mean vertical peak force per total weight (Vpk) for the hindlimb of 1.19 (sd = 0.13), consistent with those of unloaded capuchins in the laboratory and for other bipedal primates, including humans. Vertical force records suggest that capuchins, along with most nonhuman primates, maintain a relatively compliant leg during both unloaded and loaded locomotion. Like all other primates, loaded capuchins maintained laterally (outward) directed medio-lateral forces, presumably to stabilize side-to-side movements of the center of mass. Mediolateral forces suggest that at near-running speeds dynamic stability diminishes the need to generate high lateral forces. Vertical force traces exhibited a measurable impact spike at foot contact in 85% of the steps recorded. An impact spike is common in human walking and running but has not been reported in other bipedal primates. This spike in humans is thought to lead to bone and cartilage damage. The earliest biped may have experienced similar impact spikes during bipedal locomotion, requiring compensatory behaviors or anatomical features. (C) 2015 Elsevier Ltd. All rights reserved.

It is often claimed that the walking gaits of primates are unusual because, unlike most other mammals, primates appear to have higher vertical peak ground reaction forces on their hindlimbs than on their forelimbs. Many researchers have argued that this pattern of ground reaction force distribution is part of a general adaptation to arboreal locomotion. This argument is frequently used to support models of primate locomotor evolution. Unfortunately, little is known about the force distribution patterns of primates walking on arboreal supports, nor do we completely understand the mechanisms that regulate weight distribution in primates. We collected vertical peak force data for seven species of primates walking quadrupedally on instrumented terrestrial and arboreal supports. Our results show that, when walking on arboreal vs. terrestrial substrates, primates generally have lower vertical peak forces on both limbs but the difference is most extreme for the forelimb. We found that force reduction occurs primarily by decreasing forelimb and, to a lesser extent, hindlimb stiffness. As a result, on arboreal supports, primates experience significantly greater functional differentiation of the forelimb and hindlimb than on the ground. These data support long-standing theories that arboreal locomotion was a critical factor in the differentiation of the forelimbs and hindlimbs in primates. This change in functional role of the forelimb may have played a critical role in the origin of primates and facilitated the evolution of more specialized locomotor behaviors.

Analysis of the teeth, orbital, and gnathic regions of the skull, and fragmentary postcranial bones provides evidence for reconstructing a behavioral profile of Amphipithecidae: Pondaungia , Amphipithecus , Myanmarpithecus (late middle Eocene, Myanmar) and Siamopithecus (late Eocene, Thailand). At 5–8 kg, Pondaungia , Amphipithecus , and Siamopithecus are perhaps the largest known Eocene primates. The dental and mandibular anatomy suggest that larger-bodied amphipithecids were hard-object feeders. The shape of the mandibular corpus and stiffened symphysis suggest an ability to resist large internal loads during chewing and to recruit significant amounts of muscle force from both the chewing and non-chewing sides of the jaw so as to increase bite force during mastication. The large spatulate upper central incisor of Pondaungia and projecting robust canines of all the larger amphipithecids suggest that incisal food preparation was important. The molars of Siamopithecus , Amphipithecus , and Pondaungia have weak shearing crests. This, and the thick molar enamel found in Pondaungia , suggests a diet of seeds and other hard objects low in fiber. In contrast, Myanmarpithecus was smaller, about 1–2 kg; its cheek teeth suggest a frugivorous diet and do not imply seed eating. Postcranial bones (humerus, ulna, and calcaneus) of a single large amphipithecid individual from Myanmar suggest an arboreal quadrupedal locomotor style like that of howler monkeys or lorises. The humeral head is rounded, proximally oriented, and the tuberosities are low indicating an extremely mobile glenohumeral joint. The great thickness of the midshaft cortical bone of the humerus implies enhanced ability to resist bending and torsion, as seen among slow moving primate quadrupeds. The elbow joint exhibits articular features for enhanced stability in habitually flexed positions, features also commonly found in slow moving arboreal quadrupeds. The short distal load arm of the calcaneus is consistent with, but not exclusive to, slow arboreal quadrupedalism, and suggests no reliance on habitual leaping. A recently recovered talus of an amphipithecid suggests a possibly more active arboreal quadrupedalism.

Lumbar vertebral morphology has been used as an indicator of locomotor behavior in living and fossil mammals. Rigidity within the lumbar region is thought to be important for increasing overall axial rigidity during various forms of locomotion, including bridging between supports, inverted quadrupedalism, gliding, and flying. However, distinguishing between those behaviors using bony features has been challenging. This study used osteological characters of the lumbar vertebrae to attempt to develop fine-grade functional distinctions among different mammalian species in order to make more complete inferences about how the axial skeleton affects locomotor behavior in extant mammals. These same lumbar characters were measured in two extinct species for which locomotor behaviors are well known, the sloth lemurs (Palaeopropithecus and Babakotia radofilai), in order to further evaluate their locomotor behaviors. Results from a principal components analysis of seven measurements, determined to be functionally significant from previous studies, demonstrate that inverted quadrupeds in the sample are characterized by dorsoventrally short and cranio-caudally expanded spinous processes, dorsally oriented transverse processes, and mediolaterally short and dorsoventrally high vertebral bodies compared with mammals that are relatively pronograde, vertical clingers, or gliders. Antipronograde mammals, dermopterans, and chiropterans also exhibit these traits, but not to the same extent as the inverted quadrupeds. In accordance with previous studies, our data show that the sloth lemur B. radofilai groups closely with antipronograde mammals like lorises, while Palaeopropithecus groups with extant sloths. These findings suggest that Palaeopropithecus was engaged in inverted quadrupedalism at a high frequency, while B. radofilai may have engaged in a more diverse array of locomotor and positional behaviors. The osteological features used here reflect differences in lumbar mobility and suggest that axial rigidity is advantageous for suspensory locomotion and possibly flight in bats.

During the past century, many anthropologists have proposed that hominoid orthograde locomotion arose in an arboreal quadrupedal ancestor with highly mobile, low weight-bearing forelimbs. However, no quantitative data comparing kinematic and kinetic aspects of forelimb use during arboreal and terrestrial quadrupedalism have been available to evaluate such theories. In this preliminary study, a spider monkey and a baboon were videotaped in three planes while walking quadrupedally on an instrumented runway and a raised instrumented horizontal pole. Forelimb angles and substrate reaction force resultants were calculated for each animal on each substrate. The quantitative data presented here support previous models for the evolution of primate locomotion that were based on theoretical biomechanics and qualitative or anecdotal evidence. In addition, this study has revealed several previously undocumented accommodations to "arboreal" quadrupedal locomotion in these two primates. While walking on the pole, compared to travel on the ground, (1) both animals adopted a "crouched" forelimb posture, but only the spider monkey abducted its arm and ulnar deviated its hand; (2) both subjects have lower resultant forces on the forelimb due to lower absolute force magnitudes and changes in the timing of component peaks; and (3) both animals reduce and reorient transverse forces. Similar accommodations to arboreal travel by both subjects appear to be mechanical requirements of arboreal locomotion. However, differences may be due to morphological differences between the subjects, or to their divergent phylogenetic history. These results are used to explore potential explanations for the morphological differences between arboreal and terrestrial primate quadrupeds in terms of bone and joint strain and to evaluate models of primate locomotor evolution.

Since Kimura et al.'s (1979) analysis of ground reaction forces during quadrupedal walking, primates are commonly pictured as being "hindlimb driven" compared to "forelimb driven" nonprimate mammals. Hindlimb dominance in primates has subsequently been interpreted as a preadaptation to human bipedalism. However, given its considerable influence, surprisingly little data are available to support this putative contrast in limb dominance. In this reconsideration of locomotor kinetics in primates, we have collected force plate data on two chimpanzees, one orangutan, two vervet monkeys, and two cats for a range of gaits and speeds. The peak vertical forces acting on the limbs as well as the braking and propulsive impulses exerted by the limbs are examined. Forces and impulses are highly variable and change with speed, gait, and the differential use of asynchronously or asymmetrically placed limbs. Peak vertical forces increase with speed. The faster gaits (trot, gallop) have, on the average, higher forces than the walk. However, there is no major change in force magnitudes at gait transitions. The mean vertical forces are higher on the hindlimbs than on the forelimbs of the primates. This difference is most pronounced in the suspensory orangutan and least pronounced in the quadrupedal vervets. Cats, on the other hand, generate higher forelimb than hindlimb vertical forces. Although our results support the overall conclusion of Kimura et al. (1979) that peak vertical forces are relatively low on the primate forelimb, they also show some variation most probably related to locomotor mode. In the majority of primate cases, the major propulsive thrust is also generated by the hindlimbs. However, in the galloping vervets, the trailing limbs are propulsive and the leading limbs braking, no matter whether these are forelimbs or hindlimbs. A similar, although less pronounced, asymmetry between trailing and leading limbs was observed in a galloping chimpanzee. Not only are primates variable with regard to the roles of the limbs in propulsion, they are also not unique among mammals in being predominantly hindlimb driven. Our cats, as well as all other nonprimate mammals so far analysed, generate greater propulsive thrust with their hindlimbs; i.e. they are also "hindlimb driven".

The quadrupedal walking gaits of most primates can be distinguished from those of most other mammals by the presence of diagonal-sequence (DS) footfall patterns and higher peak vertical forces on the hindlimbs compared to the forelimbs. The walking gait of the woolly opossum (Caluromys philander), a highly arboreal marsupial, is also characterized by diagonal-sequence footfalls and relatively low peak forelimb forces. Among primates, three species--Callithrix, Nycticebus, and Loris--have been reported to frequently use lateral-sequence (LS) gaits and experience relatively higher peak vertical forces on the forelimbs. These patterns among primates and other mammals suggest a strong association between footfall patterns and force distribution on the limbs. However, current data for lorises are limited and the frequency of DS vs. LS walking gaits in Loris is still ambiguous. To test the hypothesis that patterns of footfalls and force distribution on the limbs are functionally linked, kinematic and kinetic data were collected simultaneously for three adult slender lorises (Loris tardigradus) walking on a 1.25 cm horizontal pole. All subjects in this study consistently used diagonal-sequence walking gaits and always had higher peak vertical forces on their forelimbs relative to their hindlimbs. These results call into question the hypothesis that a functional link exists between the presence of diagonal-sequence walking gaits and relatively higher peak vertical forces on the hindlimbs. In addition, this study tested models that explain patterns of force distribution based on limb protraction angle or limb compliance. None of the Loris subjects examined showed kinematic patterns that would support current models proposing that weight distribution can be adjusted by actively shifting weight posteriorly or by changing limb stiffness. These data reveal the complexity of adaptations to arboreal locomotion in primates and indicate that diagonal-sequence walking gaits and relatively low forelimb forces could have evolved independently.

The evolution of primates is intimately linked to their initial invasion of an arboreal environment. However, moving and foraging in this milieu creates significant mechanical challenges related to the presence of substrates differing in their size and orientation. It is widely assumed that primates are behaviorally and anatomically adapted to movement on specific substrates, but few explicit tests of this relationship in an evolutionary context have been conducted. Without direct tests of form-function relationships in living primates it is impossible to reliably infer behavior in fossil taxa. In this study, we test a hypothesis of co-variation between forelimb morphology and the type of substrates used by strepsirrhines. If associations between anatomy and substrate use exist, these can then be applied to better understand limb anatomy of extinct primates. The co-variation between each forelimb long bone and the type of substrate used was studied in a phylogenetic context. Our results show that despite the presence of significant phylogenetic signal for each long bone of the forelimb, clear support use associations are present. A strong co-variation was found between the type of substrate used and the shape of the radius, with and without taking phylogeny into account, whereas co-variation was significant for the ulna only when taking phylogeny into account. Species that use a thin branch milieu show radii that are gracile and straight and have a distal articular shape that allows for a wide range of movements. In contrast, extant species that commonly use large supports show a relatively robust and curved radius with an increased surface area available for forearm and hand muscles in pronated posture. These results, especially for the radius, support the idea that strepsirrhine primates exhibit specific skeletal adaptations associated with the supports that they habitually move on. With these robust associations in hand it will be possible to explore the same variables in extinct early primates and primate relatives and thus improve the reliability of inferences concerning substrate use in early primates.