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The Kromdraai site in South Africa has yielded numerous early hominin fossils since 1938. As a part of recent excavations within Unit P, a largely complete early hominin calcaneus (KW 6302) was discovered. Due to its role in locomotion, the calcaneus has the potential to reveal important form/function relationships. Here, we describe KW 6302 and analyze its preserved morphology relative to human and nonhuman ape calcanei, as well as calcanei attributed to Australopithecus afarensis, Australopithecus africanus, Australopithecus sediba, Homo naledi, and the Omo calcaneus (either Paranthropus or early Homo). KW 6302 calcaneal morphology is assessed using numerous quantitative metrics including linear measures, calcaneal robusticity index, relative lateral plantar process position, Achilles tendon length reconstruction, and a three-dimensional geometric morphometric sliding semilandmark analysis. KW 6302 exhibits an overall calcaneal morphology that is intermediate between humans and nonhuman apes, although closer to modern humans. KW 6302 possesses many traits that indicate it was likely well-adapted for terrestrial bipedal locomotion, including a relatively flat posterior talar facet and a large lateral plantar process that is similarly positioned to modern humans. It also retains traits that indicate that climbing may have remained a part of its locomotor repertoire, such as a relatively gracile tuber and a large peroneal trochlea. Specimens from Kromdraai have been attributed to either Paranthropus robustus or early Homo; however, there are no definitively attributed calcanei for either genus, making it difficult to taxonomically assign this specimen. KW 6302 and the Omo calcaneus, however, fall outside the range of expected variation for an extant genus, indicating that if the Omo calcaneus was Paranthropus, then KW 6302 would likely be attributed to early Homo (or vice versa).

Quantifying morphological variation is critical for conducting anatomical research. Three-dimensional geometric morphometric (3D GM) landmark analyses quantify shape using homologous Cartesian coordinates (landmarks). Setting up a high-density landmark set and placing it on all specimens, however, can be a time-consuming task. Weighted spherical harmonics (SPHARM) provides an alternative method for analyzing the shape of such objects. Here we compare sliding semilandmark and SPHARM analyses of the calcaneus of Gorilla gorilla gorilla (n = 20), Pan troglodytes troglodytes (n = 20), and Homo sapiens (n = 20) to determine whether the SPHARM and sliding semilandmark analyses capture comparable levels of shape variation. We also compare both the sliding semilandmark and SPHARM analyses to a novel combination of the two methods, here termed SPHARM-sliding. In SPHARM-sliding, the vertices of the surface models produced from the SPHARM analysis (that are the same in number and relative location) are used as the starting landmark positions for a sliding semilandmark analysis. Calcaneal shape variation quantified by all three analyses was summarized using separate principal components analyses. Results were compared using the root mean square (RMS) and maximum distance between surface models of species averages scaled (up) to centroid size created from each analysis. The average RMS was 0.23 mm between sliding semilandmark and SPHARM average surface models, 0.19 mm between SPHARM and SPHARM sliding average surface models, and 0.22 mm between sliding semilandmark and SPHARM sliding average surface models. Although results indicate that all three analyses are comparable methods for 3D shape analysis, there are advantages and disadvantages to each. While the SPHARM analysis is less time-intensive, it is unable to capture the same level of detail around the sharp edges of articular facets on average surface models as the sliding semilandmark analysis. The SPHARM analysis also does not allow for individual articular facets to be analyzed in isolation. SPHARM-sliding, however, captures the same level of detail as the sliding semilandmark analysis, and (as in the sliding semilandmark analysis) allows for the evaluation of individual portions of bone. SPHARM is a comparable method to a 3D GM analysis for small, irregularly shaped bones, such as the calcaneus, and SPHARM-sliding allows for an expedited set up process for a sliding semilandmark analysis.

Limb bone articular and diaphyseal proportions have been shown to relate to locomotor behavior in broad comparisons across catarrhines, but comparisons among phylogenetically and functionally more closely related species may be particularly useful in investigating form-function relationships that can be applied to fossil taxa. Here we compare inter- and intra-limb proportions of diaphyseal strength and articular surface area and breadth of the femur and humerus with frequencies of leaping and vertical climbing behavior in 13 cercopithecid species. Leaping frequency is highly positively correlated with femoral/humeral diaphyseal strength, moderately positively correlated with femoral/humeral articular breadth, and less highly correlated with femoral/humeral articular surface area. These results are consistent with predicted higher bending loads as well as joint reaction forces on the femora of leapers. Surface areas may show a weaker association because they also directly impact joint excursion and are thus more influenced by other aspects of locomotion, including climbing. Climbing frequency is positively correlated with humeral head articular surface area/diaphyseal strength, but weakly negatively correlated with femoral head articular surface area/diaphyseal strength. These combined trends lead to a strong negative association between climbing and femoral/humeral head surface area. Femoral/humeral diaphyseal strength and distal articular breadth are not correlated with climbing frequency. The climbing results are consistent with greater shoulder mobility in more frequent vertical climbers. The lack of such a relationship in the femur among these taxa contrasts with earlier findings for catarrhines more generally, including hominoids, and may be a result of different climbing kinematics in cercopithecoids involving less hip abduction than in hominoids. Different use of the forelimb during climbing in cercopithecoids and hominoids may also explain the lack of association between femoral/humeral diaphyseal strength and climbing in the present study, in contrast to comparisons across catarrhines more generally.

Among human and nonhuman apes, calcaneal morphology exhibits significant variation that has been related to locomotor behavior. Due to its role in weight-bearing, however, both body size and locomotion may impact calcaneal morphology. Determining how calcaneal morphologies vary as a function of body size is thus vital to understanding calcaneal functional adaptation. Here, we study calcaneus allometry and relative size in humans (n = 120) and nonhuman primates (n = 278), analyzing these relationships in light of known locomotor behaviors. Twelve linear measures and three articular facet surface areas were collected on calcaneus surface models. Body mass was estimated using femoral head superoinferior breadth. Relationships between calcaneal dimensions and estimated body mass were analyzed across the sample using phylogenetic least squares regression analyses (PGLS). Differences between humans and pooled nonhuman primates were tested using RMA ANCOVAs. Among (and within) genera residual differences from both PGLS regressions and isometry were analyzed using ANOVAs with post hoc multiple comparison tests. The relationships between all but two calcaneus dimensions and estimated body mass exhibit phylogenetic signal at the smallest taxonomic scale. This signal disappears when reanalyzed at the genus level. Calcaneal morphology varies relative to both body size and locomotor behavior. Humans have larger calcanei for estimated body mass relative to nonhuman primates as a potential adaptation for bipedalism. More terrestrial taxa exhibit longer calcaneal tubers for body mass, increasing the triceps surae lever arm. Among nonhuman great apes, more arboreal taxa have larger cuboid facet surface areas for body mass, increasing calcaneocuboid mobility.

The foot has played a prominent role in evaluating early hominin locomotion. The calcaneus, in particular, plays an important role in weight-bearing. Although the calcanei of early hominins have been previously scrutinized, a three-dimensional analysis of the entire calcaneal shape has not been conducted. Here, we investigate the relationship between external calcaneal shape and locomotion in modern Homo sapiens (n = 130), Gorilla (n = 86), Pan (n = 112), Pongo (n = 31), Papio (n = 28), and hylobatids (Hylobates, Symphalangus; n = 32). We use these results to place the calcanei attributed to Australopithecus sediba, A. africanus, A. afarensis, H. naledi, and Homo habilis/Paranthropus boisei into a locomotor context. Calcanei were scanned using either surface scanning or micro-CT and their external shape analyzed using a three-dimensional geometric morphometric sliding semilandmark analysis. Blomberg's K statistic was used to estimate phylogenetic signal in the shape data. Shape variation was summarized using a principal components analysis. Procrustes distances between all taxa as well as distances between each fossil and the average of each taxon were calculated. Blomberg's K statistic was small (K = 0.1651), indicating weak phylogenetic effects, suggesting variation is driven by factors other than phylogeny (e.g., locomotion or body size). Modern humans have a large calcaneus relative to body size and display a uniquely convex cuboid facet, facilitating a rigid midfoot for bipedalism. More arboreal great apes display relatively deeper cuboid facet pivot regions for increased midfoot mobility. Australopithecus afarensis demonstrates the most human-like calcaneus, consistent with obligate bipedalism. Homo naledi is primarily modern human-like, but with some intermediate traits, suggesting a different form of bipedalism than modern humans. Australopithecus africanus, A. sediba, and H. habilis/P. boisei calcanei all possess unique combinations of human and nonhuman ape-like morphologies, suggesting a combination of bipedal and arboreal behaviors.

Larger terrestrial mammals have generally been found to use more extended limb postures, a mechanism which maintains muscular requirements at larger sizes by improving the effective mechanical advantage (EMA) of limb musculature. Felids, however, have been documented to maintain joint angles across body sizes. If felid morphology scales isometrically, it would mean larger felids have relatively weaker muscles, compromising locomotor activities. Here, we examine the allometric relationships between the EMA of the elbow and knee extensors and body mass, finding that the EMA of the triceps brachii and quadriceps muscles scale with positive allometry. When species-specific joint angles were used rather than felid-average joint angles, EMA scales to body mass with more positive allometry. When the scaling of the muscle and ground reaction force (GRF) lever arms were investigated individually the allometric signal was lost; however, the muscle lever arms generally have allometric slope coefficients that are consistent with positive allometry, while the GRF lever arms demonstrate negative allometric slope coefficients. This suggests there are subtle alterations to limb morphology allowing different felid species to achieve an increased EMA via distinctive mechanisms. The quadriceps EMA was found to scale with sufficient positive allometry to compensate for increases in size without alteration in muscular anatomy; however, this is not the case for the triceps brachii EMA. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 302:775-784, 2019. © 2018 Wiley Periodicals, Inc.

OBJECTIVES The primate foot has been extensively investigated because of its role in weight-bearing; however, the calcaneus has been relatively understudied. Here we examine entire gorilla calcaneal external shape to understand its relationship with locomotor behavior. MATERIALS AND METHODS Calcanei of Gorilla gorilla gorilla (n = 43), Gorilla beringei graueri (n = 20), and Gorilla beringei beringei (n = 15) were surface or micro-CT scanned. External shape was analyzed through a three-dimensional geometric morphometric sliding semilandmark analysis. Semilandmarks were slid relative to an updated Procrustes average in order to minimize the bending energy of the thin plate spline interpolation function. Shape variation was summarized using principal components analysis of shape coordinates. Procrustes distances between taxa averages were calculated and resampling statistics run to test pairwise differences. Linear measures were collected and regressed against estimated body mass. RESULTS All three taxa exhibit statistically different morphologies (p

Extinct sloths represent a wide range of morphological, locomotor, and body size variation. Researchers have examined femoral neck angle in two dimensions to hypothesize locomotor behaviors in this group; however, this measure does not account for femoral neck anteversion. Here, we present a new method for quantifying femoral neck anteversion angle, in addition to femoral neck angle, to capture the 3D position of the femoral head/neck. Femora of extant (n = 21; Bradypus and Choloepus) and extinct (n = 49; Acratocnus, Megalocnus, Neocnus, and Parocnus) sloths were surface scanned and their surface models used to calculate three angles of femoral neck anteversion and femoral neck angle. Femoral neck anteversion was calculated as the angle between the femoral neck axis and the geometric axis of the femoral condyles (GA), the 35% cross section axis, and a trochanter axis. Femoral neck angle was calculated as the angle between the femoral neck and shaft axes. Genera were compared using ANOVAs with post hoc multiple comparisons for each angle. Femoral neck angle and femoral neck anteversion relative to the cross section were also analyzed. Significant differences among genera exist for all angles, (p < .001) but not all angles separate all genera. Femoral neck and anteversion angles typically yield different results, demonstrating the utility of analyzing both angles. The GA and cross section angles are highly correlated in sloths, with the exception of comparisons among Megalocnus, Parocnus, and Neocnus, suggesting morphological variation in the distal femur. While this method was applied to sloths, it has broad applicability to mammalian groups.

Relative fibular/tibial strength has been demonstrated to vary with locomotor behavior among anthropoid primates. In this study fibular/tibial strength was determined in KNM-WT 15000, a juvenile Homo erectus individual (1.5 Ma), and in OH 35, a Homo habilis (or possibly Paranthropus boisei) individual (1.8 Ma), and compared to that of adult modern humans (n = 79), chimpanzees (n = 16), gorillas (n = 16) and orangutans (n = 11). Ontogenetic changes in fibular/tibial strength were also analyzed due to KNM-WT 15000's juvenile status. Cross-sectional properties at midshaft were derived from multi-plane radiography and external contours, or CT scanning. Comparisons of log-transformed fibular/tibial polar second moment of area and anteroposterior (A-P) and mediolateral (M-L) second moments of area were carried out between extant species. Fossil deviations from each extant taxon's mean proportion were calculated in standard deviation (SD) units for that taxon. Great apes differ significantly from modern humans, with relatively stronger fibulae, particularly in the M-L plane. KNM-WT 15000 is more than 2 SD from all great apes (≥3 SD in the M-L plane) and within 1 SD of modern humans for almost all variables. This is not a result of its age, as fibular/tibial strength slightly decreases with age (i.e., becomes less like that of great apes) in humans. OH 35 falls within 1 SD of chimpanzees and orangutans for the majority of cross-sectional proportions, but more than 1 SD from humans. KNM-WT 15000 is demonstrated to be fully modern, complimenting other indications of complete terrestrial bipedality and possibly showing adaptations for endurance running. OH 35 has some human-like features; however, the relative strength of the two bones aligns the specimen with great apes, consistent with a significant degree of arboreality, in particular, vertical climbing.