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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 ofGorilla gorilla gorilla(n = 20),Pan troglodytes troglodytes(n = 20), andHomo 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.

 

The foot plays a prominent role in weight-bearing suggesting it may reflect locomotor variation. Despite the immense amount of foot research, the calcaneus has been relatively understudied. Here we analyzed the entire calcaneal shape of Gorilla gorilla gorilla (n=41), Gorilla beringei graueri (n=17) and Gorilla beringei beringei (n=8) to understand how morphology relates to locomotor behavior. Calcanei were surface scanned and external shape analyzed using a three-dimensional geometric morphometric sliding semilandmark analysis. Semilandmarks were slid to minimize the bending energy of the thin plate spline interpolation function relative to the updated Procrustes average. Generalized Procrustes Analysis was used to align landmark configurations and shape variation was summarized using a principal components analysis. Procrustes distances between species were calculated and resampling statistics were run to test for group differences. All subspecies demonstrate statistically different morphologies (p<0.005 for pairwise comparisons). G. b. graueri separates from other subspecies based on posterolateral morphology, with G. b. graueri demonstrating an elongated peroneal trochlea, and thus more bone superiorly than G. g. gorilla. Compared to G. b. beringei, G. b. graueri has less bone inferiorly near the tuberosity. Cuboid and posterior talar facet shapes correlate with arboreality. G. b. beringei (most terrestrial) has a flatter cuboid facet and a more transversely oriented/relatively smaller posterior talar facet than G. g. gorilla (most arboreal) and G. b. graueri represents an intermediate morphology. These differences demonstrate a relationship between calcaneal shape and locomotor behavior and suggest that G. b. graueri may load its foot differently from the other subspecies. This project was supported by NSF grant # BCS - 1824630. 

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.

 

A number of studies have demonstrated the ontogenetic plasticity of long bone diaphyseal structure in response to mechanical loading. Captivity should affect mechanical loading of the limbs, but whether captive apes grow differently than wild apes has been debated. Here, we compare captive and wild juvenile and adultGorillato ascertain whether growth trajectories in cross‐sectional diaphyseal shape are similar in the two environments.

A sample of young juvenile (n = 4) and adult (n = 10) captiveGorilla gorillagorillaspecimens, with known life histories, were compared with age‐matched wildG.g. gorilla(n = 62) andG. beringei beringei(n = 75) in relative anteroposterior to mediolateral bending strength of the femur, tibia, and humerus. Cross sections were obtained using peripheral quantitative CT.

Captive and wild adultG.g. gorilladiffered in bending strength ratios for all three bones, but these differences were not present in young juvenileG.g. gorilla. In comparisons across taxa, captive juvenileG.g. gorillawere more similar to wildG.g. gorillathan toG.b. beringei, while captive adultG.g. gorillawere more similar in shape toG.b. beringeiin the hind limb.

Captive and wildG. gorillafollow different ontogenetic trajectories in long bone diaphyseal shape, corresponding to environmental differences and subsequent modified locomotor behaviors. Differences related to phylogeny are most evident early in development.

 

Previous studies associate females who died in young adulthood with narrower obstetric pelvic dimensions, presumably in association with obstetric insufficiency (though this causal relationship is unresolved). In this study, we examine whether females within groups living at higher latitudes present this pattern, as high‐latitude groups have larger pelvic dimensions than groups previously examined. These patterns are compared with males. We assess whether there is evidence for younger ages‐at‐death in females to have been in response to natural selection against narrower true pelvis dimensions.

We measured 14 pelvic dimensions in 327 adults (188 females, 139 males), representing archaeological sites from mid‐latitude and high‐latitude North America. Individuals were placed into a “young” or “not young” age‐at‐death category. Latitude, sex, and age‐at‐death groups were compared using ANOVAs and scaled variance, and evidence for selection was examined withF‐tests.

Pelvic dimensions were larger in high‐latitude females and males. Females but not males who died at younger ages had smaller pelvic canals than older individuals, especially in the mediolateral inlet and anteroposterior outlet dimensions. Variance in all pelvic dimensions is equal between the two female age‐at‐death groups.

We found narrower obstetrical dimensions in the female pelvis among individuals who died at younger ages. However, statistically equivalent variances in the two female age‐at‐death groups does not support natural selection on pelvic dimensions as leading to younger ages at death. We instead argue that this difference is result of continued growth due to remodeling in the pelvis occurring in females, but not males, after early adulthood.