The relative size of the calcaneal tuber reflects heel strike plantigrady in African apes and humans.

OBJECTIVES: The positional repertoire of the human-chimpanzee last common ancestor is critical for reconstructing the evolution of bipedalism. African apes and humans share a heel strike plantigrade foot posture associated with terrestriality. Previous research has established that modern humans have a relatively large and intrinsically robust calcaneal tuber equipped to withstand heel strike forces associated with bipedal walking and running. However, it is unclear whether African apes have a relatively larger calcaneal tuber than non-heel-striking primates, and how this trait might have evolved among anthropoids. Here, I test the hypothesis that heel-striking primates have a relatively larger calcaneal tuber than non-heel-striking primates. METHODS: The comparative sample includes 331 individuals and 53 taxa representing hominoids, cercopithecoids, and platyrrhines. Evolutionary modeling was used to test for the effect of foot posture on the relative size of the calcaneal tuber in a phylogenetic framework that accounts for adaptation and inertia. Bayesian evolutionary modeling was used to identify selective regime shifts in the relative size of the calcaneal tuber among anthropoids. RESULTS: The best fitting evolutionary model was a Brownian motion model with regime-dependent trends characterized by relatively large calcaneal tubers among African apes and humans. Evolutionary modeling provided support for an evolutionary shift toward a larger calcaneal tuber at the base of the African ape and human clade. CONCLUSIONS: The results of this study support the view that African apes and humans share derived traits related to heel strike plantigrady, which implies that humans evolved from a semi-terrestrial quadrupedal ancestor.

Morphological and evolutionary insights into the keystone element of the human foot's medial longitudinal arch.

The evolution of the medial longitudinal arch (MLA) is one of the most impactful adaptations in the hominin foot that emerged with bipedalism. When and how it evolved in the human lineage is still unresolved. Complicating the issue, clinical definitions of flatfoot in living Homo sapiens have not reached a consensus. Here we digitally investigate the navicular morphology of H. sapiens (living, archaeological, and fossil), great apes, and fossil hominins and its correlation with the MLA. A distinctive navicular shape characterises living H. sapiens with adult acquired flexible flatfoot, while the congenital flexible flatfoot exhibits a 'normal' navicular shape. All H. sapiens groups differentiate from great apes independently from variations in the MLA, likely because of bipedalism. Most australopith, H. naledi, and H. floresiensis navicular shapes are closer to those of great apes, which is inconsistent with a human-like MLA and instead might suggest a certain degree of arboreality. Navicular shape of OH 8 and fossil H. sapiens falls within the normal living H. sapiens spectrum of variation of the MLA (including congenital flexible flatfoot and individuals with a well-developed MLA). At the same time, H. neanderthalensis seem to be characterised by a different expression of the MLA.

Inferring lumbar lordosis in Neandertals and other hominins.

Lumbar lordosis is a key adaptation to bipedal locomotion in the human lineage. Dorsoventral spinal curvatures enable the body's center of mass to be positioned above the hip, knee, and ankle joints, and minimize the muscular effort required for postural control and locomotion. Previous studies have suggested that Neandertals had less lordotic (ventrally convex) lumbar columns than modern humans, which contributed to historical perceptions of postural and locomotor differences between the two groups. Quantifying lower back curvature in extinct hominins is entirely reliant upon bony correlates of overall lordosis, since the latter is significantly influenced by soft tissue structures (e.g. intervertebral discs). Here, we investigate sexual dimorphism, ancestry, and lifestyle effects on lumbar vertebral body wedging and inferior articular facet angulation, two features previously shown to be significantly correlated with overall lordosis in living individuals, in a large sample of modern humans and Neandertals. Our results demonstrate significant differences between postindustrial cadaveric remains and archaeological samples of people that lived preindustrial lifestyles. We suggest these differences are related to activity and other aspects of lifestyle rather than innate population (ancestry) differences. Neandertal bony correlates of lumbar lordosis are significantly different from all human samples except preindustrial males. Therefore, although Neandertals demonstrate more bony kyphotic wedging than most modern humans, we cast doubt on proposed locomotor and postural differences between the two lineages based on inferred lumbar lordosis (or lack thereof), and we recommend future research compare fossils to modern humans from varied populations and not just recent, postindustrial samples.

Ardipithecus hand provides evidence that humans and chimpanzees evolved from an ancestor with suspensory adaptations.

The morphology and positional behavior of the last common ancestor of humans and chimpanzees are critical for understanding the evolution of bipedalism. Early 20th century anatomical research supported the view that humans evolved from a suspensory ancestor bearing some resemblance to apes. However, the hand of the 4.4-million-year-old hominin Ardipithecus ramidus purportedly provides evidence that the hominin hand was derived from a more generalized form. Here, we use morphometric and phylogenetic comparative methods to show that Ardipithecus retains suspensory adapted hand morphologies shared with chimpanzees and bonobos. We identify an evolutionary shift in hand morphology between Ardipithecus and Australopithecus that renews questions about the coevolution of hominin manipulative capabilities and obligate bipedalism initially proposed by Darwin. Overall, our results suggest that early hominins evolved from an ancestor with a varied positional repertoire including suspension and vertical climbing, directly affecting the viable range of hypotheses for the origin of our lineage.

A nearly complete foot from Dikika, Ethiopia and its implications for the ontogeny and function of Australopithecus afarensis.

The functional and evolutionary implications of primitive retentions in early hominin feet have been under debate since the discovery of Australopithecus afarensis. Ontogeny can provide insight into adult phenotypes, but juvenile early hominin foot fossils are exceptionally rare. We analyze a nearly complete, 3.32-million-year-old juvenile foot of A. afarensis (DIK-1-1f). We show that juvenile A. afarensis individuals already had many of the bipedal features found in adult specimens. However, they also had medial cuneiform traits associated with increased hallucal mobility and a more gracile calcaneal tuber, which is unexpected on the basis of known adult morphologies. Selection for traits functionally associated with juvenile pedal grasping may provide a new perspective on their retention in the more terrestrial adult A. afarensis.

Reevaluating the functional implications of Australopithecus afarensis navicular morphology.

The longitudinal arch is a unique characteristic of the human foot, yet the timing and pattern of its evolution remain controversial, in part due to the disagreement among researchers over which skeletal traits are the best indicators of its presence or absence. The small size of the human navicular tuberosity has previously been linked to the presence of a longitudinal arch, implying that the large tuberosity of early hominins such as Australopithecus afarensis reflects a flat foot. However, this hypothesis is at odds with other evidence of pedal form and function, such as metatarsal, tarsal, and footprint morphology, which show that a longitudinal arch was probably present in A. afarensis. This study reevaluates the morphometric affinities of the A. afarensis naviculars among other Plio-Pleistocene fossil hominins and anthropoid primates (N = 170). Multivariate cluster analyses show that all fossil hominin naviculars, including those attributed to A. afarensis, are most similar to modern humans. A measure of navicular tuberosity size quantified as the ratio of the tuberosity volume to the surface area of the talar facet shows that Ateles has the largest navicular tuberosity among the anthropoid sample and that there is no difference between highly arboreal and terrestrial taxa in this metric (e.g., Hylobates and Gorilla beringei). Instead, a relatively large navicular tuberosity may reflect the development of leg musculature associated with ankle plantarflexion. The functional inferences derived from the morphology of the A. afarensis naviculars are consistent with the morphology of the Laetoli footprints.

Conarticular congruence of the hominoid subtalar joint complex with implications for joint function in Plio-Pleistocene hominins.

OBJECTIVE: The purpose of this study is to test the hypothesis that conarticular surfaces areas and curvatures are correlates of mobility at the hominoid talocalcaneal and talonavicular joints. MATERIALS AND METHODS: Articular surface areas and curvatures of the talonavicular, anterior talocalcaneal, and posterior talocalcaneal joints were quantified using a total of 425 three-dimensional surface models of extant hominoid and fossil hominin tali, calcanei, and naviculars. Quadric surface fitting was used to calculate curvatures, pairwise comparisons were used to evaluate statistical differences between taxa, and regression was used to test for the effects of allometry. RESULTS: Pairwise comparisons show that the distributions of values for joint curvature indices follow the predicted arboreal-terrestrial morphocline in hominoid primates with no effect of body mass (PGLS p > 0.05). OH 8 (Homo habilis) and LB 1 (Homo floresiensis) can be accommodated within the range of human variation for the talonavicular joint, whereas MH2 (Australopithecus sediba) falls within the ranges of variation for Pan troglodytes and Gorilla gorilla in measures of posterior talocalcaneal joint congruity. CONCLUSIONS: Joint curvature indices are better discriminators than joint surface area indices, which may reflect a greater contribution of rotation, rather than translation, to joint movement in plantigrade taxa due to discrepancies in conarticular congruence and the "convex-concave" rule. The pattern of joint congruence in Au. sediba contributes to other data on the foot and ankle suggesting that the lateral side of the foot was more mobile than the medial side, which is consistent with suggestions of increased medial weight transfer associated with hyperpronation. Am J Phys Anthropol 160:446-457, 2016. © 2016 Wiley Periodicals, Inc.

The subtalar joint complex of Australopithecus sediba.

The hominin talus has figured prominently in previous studies of the functional morphology of the talocrural joint, but the talocalcaneal and talonavicular joints have received comparatively less attention despite their functional importance as components of the subtalar joint complex. An associated complete talus and calcaneus attributed to the Malapa Hominin 2 (MH2) individual of Australopithecus sediba offers the opportunity to evaluate the subtalar joint complex in an early hominin. Furthermore, detailed morphological comparisons of A. sediba to other fossil hominins such as Australopithecus africanus have not yet been conducted. Here I quantify joint curvatures and angular measurements among extant hominoids and fossil hominins to evaluate the functional morphology of the subtalar joint complex of A. sediba. Australopithecus sediba uniquely combines talocalcaneal joint morphology indicative of mobility with specializations of the talonavicular joint that provide medial midtarsal stabilization. Multivariate analyses of talus and calcaneus variables show that A. sediba is most similar to extant gorillas in the morphology of the subtalar joint complex. In contrast, other hominins, such as OH 8, are more similar to modern humans. The morphological similarity between MH2 (U.W. 88-98/99) and specimens from Sterkfontein, Member 4 (StW 88, StW 102, StW 352) in morphologies of the talonavicular and talocalcaneal joints suggests that A. sediba may have possessed a foot that was functionally similar to that of A. africanus. This combination of morphologies in the A. sediba foot is probably derived among hominins and suggests that arboreality may have been adaptively significant for southern African Australopithecus.

Rearfoot posture of Australopithecus sediba and the evolution of the hominin longitudinal arch.

The longitudinal arch is one of the hallmarks of the human foot but its evolutionary history remains controversial due to the fragmentary nature of the fossil record. In modern humans, the presence of a longitudinal arch is reflected in the angular relationships among the major surfaces of the human talus and calcaneus complex, which is also known as the rearfoot. A complete talus and calcaneus of Australopithecus sediba provide the opportunity to evaluate rearfoot posture in an early hominin for the first time. Here I show that A. sediba is indistinguishable from extant African apes in the angular configuration of its rearfoot, which strongly suggests that it lacked a longitudinal arch. Inferences made from isolated fossils support the hypothesis that Australopithecus afarensis possessed an arched foot. However, tali attributed to temporally younger taxa like Australopithecus africanus and Homo floresiensis are more similar to those of A. sediba. The inferred absence of a longitudinal arch in A. sediba would be biomechanically consistent with prior suggestions of increased midtarsal mobility in this taxon. The morphological patterns in talus and calcaneus angular relationships among fossil hominins suggest that there was diversity in traits associated with the longitudinal arch in the Plio-Pleistocene.

Calcaneal robusticity in Plio-Pleistocene hominins: implications for locomotor diversity and phylogeny.

A key pedal adaptation to bipedality is a relatively large, weight-bearing calcaneus. The earliest evidence for a human-like, robust calcaneus is at 3.2 Ma in Australopithecus afarensis (A.L. 333-8, A.L. 333-55, A.L. 333-37) from Hadar, Ethiopia. Australopithecus sediba at 1.98 Ma from Malapa, South Africa displays a unique combination of primitive australopith features and more derived Homo-like features, but surprisingly is characterized by a gracile, chimpanzee-like calcaneus. The differences in calcaneal morphology suggest that these taxa differed in the frequency of arboreality and in the manner of foot function during terrestrial bipedal locomotion. This study examines calcaneal morphology in extant hominids (i.e., great apes and humans; N = 95) and fossil hominins (N = 5) to better understand the evolutionary development of calcaneal robusticity in early hominins. In particular, this study focuses on two additional fossil hominin calcanei that have not figured prominently in previous discussions of calcaneal robusticity: StW 352 and Omo 33-74-896. A measure of calcaneal robusticity was quantified as the ratio of calcaneal tuber cross-sectional area to calcaneal tuber length, which significantly differs between humans and non-humans using a sequential Bonferroni alpha adjustment for multiple comparisons. Additional multivariate analyses using Mosimann shape variables show that StW 352 and Omo 33-74-896 are more similar to Au. sediba in calcaneal tuber morphology than to Au. afarensis, suggesting that the latter taxon is better adapted for terrestrial bipedalism than at least some later species of Australopithecus. This finding implies the possibility of several complex evolutionary scenarios involving either multiple reversals in postcranial morphology in Australopithecus or the independent acquisition of adaptations to terrestrial bipedalism in Au. afarensis and Homo.