Reflecting these differences in foot–ground contact between humans and African great apes during locomotion, the calcaneus morphology differs largely among species. Old world monkeys, such as Japanese macaques, usually touch down with the fore- and midfoot but do not walk with a heel strike 1, 8, 9. African great apes also have plantigrade feet and can walk with a heel strike 1, 3, 4, 5, 6, 7, but heel and midfoot often contact the ground at the same time 6, and the heel strike in African great apes is not as prominent as that in humans. The human foot possesses a calcaneus, the tuberosity of which points posteriorly and inferiorly, allowing a prominent heel strike during bipedal walking 1, 2. Such information might be useful for inferring foot functions and reconstructing the locomotion of fossil hominoids and hominids. These interspecific differences possibly reflect the functional adaptation of the calcaneus to locomotor behavior in great apes. However, the most terrestrial gorillas exhibited longer lever arm of the triceps surae muscle, larger peroneal trochlea, more concave plantar surface, more inverted calcaneal tuberosity, more everted cuboid articular surface, and more prominent plantar process than the orangutans and chimpanzees. The most arboreal orangutans possessed comparatively more slender calcaneal tuberosity and deeper pivot region of the cuboid articular surface than chimpanzees and gorillas. Clear interspecific differences in calcaneal morphology were extracted, corresponding to the degree of arboreality of the three species. A total of 556 landmarks and semilandmarks were placed on the calcaneal surface to calculate the principal components of shape variations among specimens. ![]() This study aims to clarify variations in calcaneal morphology among great apes based on three-dimensional geometric morphometrics. However, morphological variations in the calcaneus of the great apes (chimpanzees, gorillas, and orangutans) have not been sufficiently studied. Investigating the morphological differences of the calcaneus in humans and great apes is crucial for reconstructing locomotor repertories of fossil hominins.
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