How Mammals Run

The stiff human foot enables an efficient push-off when walking or running, and was critical for the evolution of bipedalism1–6. The uniquely arched morphology of the human midfoot is thought to stiffen it5–9, whereas other primates have flat feet that bend severely in the midfoot7,10,11. However, the relationship between midfoot geometry and stiffness remains debated in foot biomechanics12,13, podiatry14,15 and palaeontology4–6. These debates centre on the medial longitudinal arch5,6 and have not considered whether stiffness is affected by the second, transverse tarsal arch of the human foot16. Here we show that the transverse tarsal arch, acting through the inter-metatarsal tissues, is responsible for more than 40% of the longitudinal stiffness of the foot. The underlying principle resembles a floppy currency note that stiffens considerably when it curls transversally. We derive a dimensionless curvature parameter that governs the stiffness contribution of the transverse tarsal arch, demonstrate its predictive power using mechanical models of the foot and find its skeletal correlate in hominin feet. In the foot, the material properties of the inter-metatarsal tissues and the mobility of the metatarsals may additionally influence the longitudinal stiffness of the foot and thus the curvature–stiffness relationship of the transverse tarsal arch. By analysing fossils, we track the evolution of the curvature parameter among extinct hominins and show that a human-like transverse arch was a key step in the evolution of human bipedalism that predates the genus Homo by at least 1.5 million years. This renewed understanding of the foot may improve the clinical treatment of flatfoot disorders, the design of robotic feet and the study of foot function in locomotion. The transverse tarsal arch, acting through the inter-metatarsal tissues, is important for the longitudinal stiffness of the foot and its appearance is a key step in the evolution of human bipedalism.

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Stiffness of the human foot and evolution of the transverse arch.

Bale monkeys (Chlorocebus djamdjamensis) are little-known primates endemic to the forests of the Bale Massif and Hagere Selam regions of Ethiopia. From August 2007 to May 2008, we conducted the first ever study of the species’ behavior and ecology, focusing in particular on its diet, activity patterns, and ranging ecology in the Odobullu Forest. We studied 2 neighboring groups (group A: 55–60 members; group B: 46–50 members) and conducted behavioral scan samples on the first 2–5 individuals sighted at 15-min intervals. Feeding accounted for 65.7% of the activity budget, followed by moving (14.4%), resting (10.7%), social (7.1%), and other behaviors (2.4%). Overall diet during the study was dominated by young leaves (80.2%), though subjects also ate fruits (9.6%), flowers (3.1%), animal prey (2.3%), shoots (1.5%), stems (1.4%), mature leaves (1.1%), and roots (0.9%). Bale monkeys consumed only 11 plant species; of these, the top 5 species accounted for 94.3% of their diet. The top food item, bamboo (Arundinaria alpina), was responsible for a remarkable 76.7% of their diet, with most (95.2%) of the bamboo consumption consisting of young leaves. Mean daily path length for the study groups was 928 m and mean (100% minimum convex polygon) home range size was 15.2 ha. Though we are cautious in drawing conclusions from only 2 groups, the larger group traveled further per day and occupied a larger home range, patterns suggesting scramble competition may be occurring in Bale monkey groups at Odobullu. The dietary specialization of Bale monkeys on bamboo makes them unique among Chlorocebus spp. and suggests an intriguing ecological convergence with the golden monkeys (Cercopithecus mitis kandti) of Uganda and bamboo lemurs (Hapalemur spp.) of Madagascar. Their narrow ecological niche, limited geographic distribution, and bamboo harvesting by local people for commercial purposes place Bale monkeys at risk of extinction. To ensure the long-term survival of Bale monkeys, appropriate management action should be taken to conserve the species and the bamboo forests upon which it depends.

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Diet, Activity Patterns, and Ranging Ecology of the Bale Monkey (Chlorocebus djamdjamensis) in Odobullu Forest, Ethiopia

Previous researchers found positive scaling of body size and sexual size dimorphism (SSD) in primates, known as Rensch's rule. The pattern is present in Haplorhini, but absent in Strepsirhini. I found that positive evolutionary correlations between size and SSD drive positive scaling relationships within Haplorhini as a whole and Platyrrhini, Cercopithecinae, Colobinae, and Hominoidea individually at the generic level and higher, but that evolutionary correlations within genera in these clades are often nonsignificant or negative. I suggest that positive evolutionary correlations result from greater change in male than in female size, usually because of sexual selection acting on polygynous populations. I suggest that negative evolutionary correlations result from greater change in female size, owing to either natural selection or, in Callitrichidae, sexual selection acting on polyandrous populations. The high incidence of negative evolutionary correlations within Haplorhini suggests a relatively large influence of natural selection on SSD, at least with regard to differences in SSD between congeners. I propose two possible explanations for the difference in intrageneric and supergeneric evolutionary patterns: 1) natural selection is a relatively weak force for modifying SSD and has a noticeable effect only when one compares related species experiencing similar levels of sexual selection, and 2) natural selection is a relatively strong force for modifying SSD but is less likely than sexual selection to affect higher level taxonomic comparisons noticeably because of the cumulative effect over time of marginal differences in mortality rates of these two types of selection. I discuss types of data required to test these explanations and implications for reconstructing fossil behavior.

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Scaling of Size and Dimorphism in Primates II: Macroevolution

This paper describes postcranial remains pertaining to the endemic xenotrichin callicebines of the Greater Antilles, all of which are extinct: Xenothrix mcgregori (Jamaica), Antillothrix bernensis (Hispaniola), and Paralouatta varonai and P. marianae (Cuba). These monkeys differed considerably in body size and inferred locomotor behavior. Xenothrix and Antillothrix are estimated to have weighed 2–5 kg, which is well within the middle range of body sizes found in extant South American monkeys, but Paralouatta (~9–10 kg) would have been nearly as large as the largest living platyrrhines. In line with previous studies, we interpret Xenothrix mcgregori as a rather short-limbed, slow-moving arboreal quadruped possessing some unusual features not otherwise seen in platyrrhines (e.g., adductor process or “fourth trochanter” of the femur). Its closest locomotor analog among living primates remains uncertain. Paralouatta varonai also exhibits features not seen in other platyrrhines, but in this case there...

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Postcranial Remains of the Extinct Monkeys of the Greater Antilles, with Evidence for Semiterrestriality in Paralouatta1

Body weight and length, chest girth, and seven postcranial limb segment lengths are compared between two guenon species, Chlorocebus (Cercopithecus) aethiops (vervets) and Cercopithecus mitis (blue monkeys), exhibiting different habitual locomotor preferences. The subjects, all adults, were wild caught for a non-related research project (Turner et al. [1986] Genetic and morphological studies on two species of Kenyan monkeys, C. aethiops and C. mitis. In: Else JG, Lee PC, editors. Primate evolution, proceedings of the Xth International Congress of Primatology, Cambridge. London). The morphological results are interpreted within the context of previously published observations of primate locomotion and social organization. The sample is unique in that the body weight of each individual is known, allowing the effects of body-size scaling to be assessed in interspecific and intersexual comparisons. C. mitis has a significantly (P < 0.05) greater body weight and trunk length than C. aethiops. A shorter trunk may function to reduce spinal flexibility for ground-running in the latter. Proximal limb segments (arm and thigh) are significantly greater in C. mitis, reflecting known adaptations to committed arboreal quadrupedal locomotion. By contrast, relative distal limb segments (forearm, crus, and foot) are significantly longer in C. aethiops, concordant with a locomotor repertoire that includes substantial terrestrial quadrupedalism, in addition to arboreal agility, and also the requisite transition between ground and canopy. Although normally associated with arboreal monkeys, greater relative tail length occurs in the more terrestrial vervets. However, because vervets exploit both arboreal and terrestrial habitats, a longer tail may compensate for diminished balance during arboreal quadrupedalism resulting from the greater "brachial" and "crural" indices that enhance their ground quadrupedalism. Most interspecific differences in body proportions are explicable by differences in locomotor modalities. Some results, however, contradict commonly held "tenets" that relate body size and morphology exclusively to locomotion. Generally associated with terrestriality, sexual dimorphism (male/female) is greater in the more arboreal blue monkeys. A more intense, seasonal mating competition may account for this incongruity.

C.S. Mott

Papers

Comparative postcranial body shape and locomotion in Chlorocebus aethiops and Cercopithecus mitis.