It has been shown that gibbons are able to brachiate with very low mechanical costs. The conversion of muscle activity into smooth, purposeful movement of the limb depends on the morphometry of muscles and their mechanical action on the skeleton. Despite the gibbon's reputation for excellence in brachiation, little information is available regarding either its gross musculoskeletal anatomy or its more detailed muscle-tendon architecture. We provide quantitative anatomical data on the muscle-tendon architecture (muscle mass, physiological cross-sectional area, fascicle length and tendon length) of the forelimb of four gibbon species, collected by detailed dissections of unfixed cadavers. Data are compared between different gibbon species and with similar published data of non-brachiating primates such as macaques, chimpanzees and humans. No quantitative differences are found between the studied gibbon species. Both their forelimb anatomy and muscle dimensions are comparable when normalized to the same body mass. Gibbons have shoulder flexors, extensors, rotator muscles and elbow flexors with a high power or work-generating capacity and their wrist flexors have a high force-generating capacity. Compared with other primates, the elbow flexors of gibbons are particularly powerful, suggesting that these muscles are particularly important for a brachiating lifestyle. Based on this anatomical study, the shoulder flexors, extensors, rotator muscles, elbow flexors and wrist flexors are expected to contribute the most to brachiation.

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We collected high-resolution plantar pressure distributions of seven bonobos during terrestrial bipedal and quadrupedal locomotion (N = 146). Functional foot length, degree of hallux abduction, and total contact time were determined, and plots, showing pressure as a function of time for six different foot regions, were generated. We also studied five adult humans for comparison (N = 13). Both locomotion types of the bonobo show a large variation in plantar pressure distributions, which could be due to the interference of instantaneous behavior with locomotion and differences in walking speed and body dimensions. The heel and the lateral midfoot typically touch down simultaneously at initial ground contact in bipedal and quadrupedal walking of bonobos, in contrast with the typical heel-strike of human bipedalism. The center of pressure follows a curved course during quadrupedalism, as a consequence of the medial weight transfer during mid-stance. Bipedal locomotion of bonobos is characterized by a more plantar positioning of the feet and by a shorter contact time than during quadrupedal walking, according to a smaller stride and step length at a higher frequency. We observed a varus position of the foot with an abducted hallux, which likely possesses an important sustaining and stabilizing function during terrestrial locomotion.

It is commonly held that the major functional features of the human foot (e.g. a functional longitudinal medial arch, lateral to medial force transfer and hallucal (big-toe) push-off ) appear only in the last 2 Myr, but functional interpretations of footbones and footprints of early human ancestors (hominins) prior to 2 million years ago (Mya) remain contradictory. Pixel-wise topographical statistical analysis of Laetoli footprint morphology, compared with results from experimental studies of footprint formation; foot-pressure measurements in bipedalism of humans and non-human great apes; and computer simulation techniques, indicate that most of these functional features were already present, albeit less strongly expressed than in ourselves, in the maker of the Laetoli G-1 footprint trail, 3.66 Mya. This finding provides strong support to those previous studies which have interpreted the G-1 prints as generally modern in aspect.

Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee and ankle joints in the orang-utan, gibbon, gorilla (Eastern and Western lowland) and bonobo . Moment arms varied with joint motion and were generally longer in proximal limb muscles than distal limb muscles. The shape of the moment arm curves (i.e. the plots of moment arm against joint angle) differed in different hindlimb muscles and in the same muscle in different subjects (both in the same and in different ape species). Most moment arms increased with increasing joint flexion, a finding which may be understood in the context of the employment of flexed postures by most nonhuman apes (except orang-utans) during both terrestrial and arboreal locomotion. When compared with humans, non-human great apes tended to have muscles better designed for moving the joints through large ranges. This was particularly true of the pedal digital flexors in orang-utans. In gibbons, the only lesser ape studied here, many of the moment arms measured were relatively short compared with those of great apes. This study was performed on a small sample of apes and thus differences noted here warrant further investigation in larger populations.

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SUMMARYThe ostrich is highly specialized in terrestrial locomotion and is the only extant bird that is both didactyl and exhibits a permanently elevated metatarsophalangeal joint. This extreme degree of digitigrady provides an excellent opportunity for the study of phalangeal adaptation towards fast, sustained bipedal locomotion. Data were gathered in a semi-natural setting with hand-raised, cooperative specimens. Dynamic pressure distribution, centre of pressure (CoP) trajectory and the positional interrelationship of the toes during stance phase were investigated using pedobarography. Walking and running trials shared a Jshaped CoP trajectory with greater localization of CoP origin as speed increased. Slight variations of 4th toe position in walking affect CoP origin and modulation of 4th toe pressure on the substrate allows correction of balance, primarily at the beginning of stance phase at lower speeds. Load distribution patterns differed significantly between slow and fast trials. In walking, the 3rd and particularly the 4th toe exhibited notable variation in load distribution with minor claw participation only at push-off. Running trials yielded a distinctly triangular load distribution pattern defined by the 4th toe tip, the proximal part of the 3rd toe and the claw tip, with the sharp point of the claw providing an essential traction element at push-off. Consistency of CoP trajectory and load distribution at higher speeds arises from dynamic stability effects and may also reflect stringent limitations to degrees of freedom in hindlimb joint articulation that contribute to locomotor efficiency. This novel research could aid in the reconstruction of theropod locomotor modes and offers a systemic approach for future avian pedobarographic investigations.

This paper illustrates how simple mechanical models based on morphological, ethological, ecological and phylogenetic data can add to discussions in evolutionary biology. Bipedal locomotion has evolved on numerous occasions in lizards. Traits that appear repeatedly in independent evolutionary lines are often considered adaptive, but the exact advantages of bipedal locomotion in lizards remain debated. Earlier claims that bipedalism would increase maximal running speed or would be energetically advantageous have been questioned. Here, we use 'whole body' mechanical modelling to provide an alternative solution to the riddle. The starting point is the intermittent running style combined with the need for a high manoeuvrability characterizing many small lizard species. Manoeuvrability benefits from a caudal shift of the centre of mass of the body (body-COM), because forces to change the heading and to align the body to this new heading do not conflict with each other. The caudally situated body-COM, however, might result in a lift of the front part of the body when accelerating (intermittent style), thus resulting in bipedal running bouts. Based on a momentum-impulse approach the effect of acceleration is quantified for a mechanical model, a virtual lizard (three segments) based on the morphometrics of Acanthodactylus erythrurus (a small lacertid lizard). Biologically relevant input (dimensions, inertial properties, step cycle information, etc.) results in an important lift of the front part of the body and observable distances passively covered bipedally as a consequence of the acceleration. In this way, no functional explanation of the phenomenon of lizard bipedalism is required and bipedalism can probably be considered non-adaptive in many cases. This does not exclude, however, some species that may have turned this consequence to their benefit. For instance, instantaneous manipulation of the position of the centre of the body-COM allows stable, persisting bipedal running. Once this was achieved, the bipedal spandrel could be exploited further.