Morphological integration in the hominid midfoot.

The calculation of morphological integration across living apes and humans may provide important insights into the potential influence of integration on evolutionary trajectories in the hominid lineage. Here, we quantify magnitudes of morphological integration among and within elements of the midfoot in great apes and humans to examine the link between locomotor differences and trait covariance. We test the hypothesis that the medial elements of the great ape foot are less morphologically integrated with one another compared to humans based on their abducted halluces, and aim to determine how adaptations for midfoot mobility/stiffness and locomotor specialization influence magnitudes of morphological integration. The study sample is composed of all cuneiforms, the navicular, the cuboid, and metatarsals 1-5 of Homo sapiens (n = 80), Pan troglodytes (n = 63), Gorilla gorilla (n = 39), and Pongo sp. (n = 41). Morphological integration was quantified using the integration coefficient of variation of interlandmark distances organized into sets of a priori-defined modules. Magnitudes of integration across these modules were then compared against sets of random traits from the whole midfoot. Results show that all nonhuman apes have less integrated medial elements, whereas humans have highly integrated medial elements, suggesting a link between hallucal abduction and reduced levels of morphological integration. However, we find considerable variation in magnitudes of morphological integration across metatarsals 2-5, the intermediate and lateral cuneiform, the cuboid, and navicular, emphasizing the influence of functional and nonfunctional factors in magnitudes of integration. Lastly, we find that humans and orangutans show the lowest overall magnitudes of integration in the midfoot, which may be related to their highly specialized functions, and suggest a link between strong diversifying selection and reduced magnitudes of morphological integration.

Mandibular shape variation in mainland and insular hylobatids.

Although hylobatids are the most speciose of the living apes, their morphological interspecies and intraspecies variation remains poorly understood. Here, we assess mandibular shape variation in two species of Hylobates, white-handed (Hylobates lar) and black-handed (Hylobates agilis) gibbons. Using 71 three-dimensional landmarks to quantify mandibular shape, interspecies and intraspecies variation and geographic patterns of mandibular shape are examined in a mixed sex sample of adult H. lar and H. agilis through generalized Procrustes analysis, Procrustes analysis of variance, and principal components analysis. We find that relative to H. agilis, H. lar exhibits a higher amount of variation in mandibular shape. Both species demonstrate similar allometric patterns in mandibular shape. We also highlight a geographic pattern in mandibular shape variation. Compared to mainland hylobatids, insular hylobatids have relatively lower, more posteriorly oriented, and anteroposteriorly wider mandibular condyles, with an increased distance between the condyles and the coronoid processes. This geographic pattern could reflect differences in functional demands on the mandible during mastication and/or could be driven by factors often associated with evolutionary pressures of island populations relative to mainland populations. The findings of this study highlight how little is known about Hylobates morphological variation and how important this is for using Hylobates to help interpret the primate fossil record. Understanding interspecific and intraspecific variation in extant primates is vital to interpreting variation in the primate fossil record.

First metatarsal trabecular bone structure in extant hominoids and Swartkrans hominins.

Changes in first metatarsal (MT1) morphology within the hominin clade are crucial for reconstructing the evolution of a forefoot adapted for human-like gait. Studies of the external morphology of the MT1 in humans, non-human apes, and fossil hominins have documented changes in its robusticity, epiphyseal shape and its articulation with the medial cuneiform. Here, we test whether trabecular structure in the MT1 reflects different loading patterns in the forefoot across extant large apes and humans, and within this comparative context, infer locomotor behavior in two fossil hominins from Swartkrans, South Africa. Microtomographic scans were collected from the MT1 of Pongo sp. (n = 6), Gorilla gorilla (n = 10), Pan troglodytes (n = 10), Homo sapiens (n = 11), as well as SKX 5017 (Paranthropus robustus), and SK 1813 (Hominin gen. sp. indet.). Trabecular structure was quantified within the head and base using a 'whole-epiphysis' approach with medtool 4.2. We found that modern humans displayed relatively higher bone volume fraction (BV/TV) in the dorsal region of each epiphysis and a higher overall degree of anisotropy (DA), whereas great apes showed higher BV/TV in the plantar regions, reflecting dorsiflexion at the metatarsophalangeal (MTP) joint in the former and plantarflexion in the latter. Both fossils displayed low DA, with SKX 5017 showing a hyper-dorsal concentration of trabecular bone in the head (similar to humans), while SK 1813 showed a more central trabecular distribution not seen in either humans or non-human apes. Additionally, we found differences between non-human apes, modern humans, and the fossil taxa in trabecular spacing (Tb.Sp.), number (Tb.N.), and thickness (Tb.th.). While low DA in both fossils suggests increased mobility of the MT1, differences in their trabecular distributions could indicate variable locomotion in these Pleistocene hominins (recognizing that the juvenile status of SK 1813 is a potential confounding factor). In particular, evidence for consistent loading in hyper-dorsiflexion in SKX 5017 would suggest locomotor behaviors beyond human-like toe off during terrestrial locomotion.