New insights into patterns of integration in the femur and pelvis among catarrhines.

OBJECTIVES: Integration reflects the level of coordinated variation of the phenotype. The integration of postcranial elements can be studied from a functional perspective, especially with regards to locomotion. This study investigates the link between locomotion, femoral structural properties, and femur-pelvis complex morphology. MATERIALS AND METHODS: We measured (1) morphological integration between femoral and pelvic morphologies using geometric morphometrics, and (2) covariation between femoral/pelvic morphologies and femoral diaphyseal cross-sectional properties, which we defined as morpho-structural integration. Morphological and morpho-structural integration patterns were measured among humans (n = 19), chimpanzees and bonobos (n = 16), and baboons (n = 14), whose locomotion are distinct. RESULTS: Baboons show the highest magnitude of morphological integration and the lowest of morpho-structural integration. Chimpanzees and bonobos show intermediate magnitude of morphological and morpho-structural integration. Yet, body size seems to have a considerable influence on both integration patterns, limiting the interpretations. Finally, humans present the lowest morphological integration and the highest morpho-structural integration between femoral morphology and structural properties but not between pelvic morphology and femur. DISCUSSION: Morphological and morpho-structural integration depict distinct strategies among the samples. A strong morphological integration among baboon's femur-pelvis module might highlight evidence for long-term adaptation to quadrupedalism. In humans, it is likely that distinct selective pressures associated with the respective function of the pelvis and the femur tend to decrease morphological integration. Conversely, high mechanical loading on the hindlimbs during bipedal locomotion might result in specific combination of structural and morphological features within the femur.

A comparative study of muscle activity and synergies during walking in baboons and humans.

Bipedal locomotion was a major functional change during hominin evolution, yet, our understanding of this gradual and complex process remains strongly debated. Based on fossil discoveries, it is possible to address functional hypotheses related to bipedal anatomy, however, motor control remains intangible with this approach. Using comparative models which occasionally walk bipedally has proved to be relevant to shed light on the evolutionary transition toward habitual bipedalism. Here, we explored the organization of the neuromuscular control using surface electromyography (sEMG) for six extrinsic muscles in two baboon individuals when they walk quadrupedally and bipedally on the ground. We compared their muscular coordination to five human subjects walking bipedally. We extracted muscle synergies from the sEMG envelopes using the non-negative matrix factorization algorithm which allows decomposing the sEMG data in the linear combination of two non-negative matrixes (muscle weight vectors and activation coefficients). We calculated different parameters to estimate the complexity of the sEMG signals, the duration of the activation of the synergies, and the generalizability of the muscle synergy model across species and walking conditions. We found that the motor control strategy is less complex in baboons when they walk bipedally, with an increased muscular activity and muscle coactivation. When comparing the baboon bipedal and quadrupedal pattern of walking to human bipedalism, we observed that the baboon bipedal pattern of walking is closer to human bipedalism for both baboons, although substantial differences remain. Overall, our findings show that the muscle activity of a non-adapted biped effectively fulfills the basic mechanical requirements (propulsion and balance) for walking bipedally, but substantial refinements are possible to optimize the efficiency of bipedal locomotion. In the evolutionary context of an expanding reliance on bipedal behaviors, even minor morphological alterations, reducing muscle coactivation, could have faced strong selection pressure, ultimately driving bipedal evolution in hominins.

Predicting primate tongue morphology based on geometrical skull matching. A first step towards an application on fossil hominins.

As part of a long-term research project aiming at generating a biomechanical model of a fossil human tongue from a carefully designed 3D Finite Element mesh of a living human tongue, we present a computer-based method that optimally registers 3D CT images of the head and neck of the living human into similar images of another primate. We quantitatively evaluate the method on a baboon. The method generates a geometric deformation field which is used to build up a 3D Finite Element mesh of the baboon tongue. In order to assess the method's ability to generate a realistic tongue from bony structure information alone, as would be the case for fossil humans, its performance is evaluated and compared under two conditions in which different anatomical information is available: (1) combined information from soft-tissue and bony structures; (2) information from bony structures alone. An Uncertainty Quantification method is used to evaluate the sensitivity of the transformation to two crucial parameters, namely the resolution of the transformation grid and the weight of a smoothness constraint applied to the transformation, and to determine the best possible meshes. In both conditions the baboon tongue morphology is realistically predicted, evidencing that bony structures alone provide enough relevant information to generate soft tissue.

From quadrupedal to bipedal walking 'on the fly': the mechanics of dynamical mode transition in primates.

We investigated how baboons transition from quadrupedal to bipedal walking without any significant interruption in their forward movement (i.e. transition 'on the fly'). Building on basic mechanical principles (momentum only changes when external forces/moments act on the body), insights into possible strategies for such a dynamical mode transition are provided and applied first to the recorded planar kinematics of an example walking sequence (including several continuous quadrupedal, transition and subsequent bipedal steps). Body dynamics are calculated from the kinematics. The strategy used in this worked example boils down to: crouch the hind parts and sprint them underneath the rising body centre of mass. Forward accelerations are not in play. Key characteristics of this transition strategy were extracted: progression speed, hip height, step duration (frequency), foot positioning at touchdown with respect to the hip and the body centre of mass (BCoM), and congruity between the moments of the ground reaction force about the BCoM and the rate of change of the total angular moment. Statistical analyses across the full sample (15 transitions of 10 individuals) confirm this strategy is always used and is shared across individuals. Finally, the costs (in J kg-1 m-1) linked to on the fly transitions were estimated. The costs are approximately double those of both the preceding quadrupedal and subsequent bipedal walking. Given the short duration of the transition as such (<1 s), it is argued that the energetic costs to change walking posture on the fly are negligible when considered in the context of the locomotor repertoire.

Development of bipedal walking in olive baboons, Papio anubis: A kinematic analysis.

OBJECTIVE: Although extant nonhuman primates are not habitual bipeds, they are able to walk bipedally from an early age. In humans, children improve their walking skills through developmental processes and learning experience. In nonhuman primates, infants do not routinely experience bipedalism and their musculoskeletal system gradually specializes for other locomotor modes. The aim of this study is to explore the development of occasional bipedal walking in olive baboon and to test whether the postural adjustments change with age. MATERIALS AND METHODS: We collected kinematics and spatiotemporal parameters of bipedal gait in an ontogenetic sample of 24 baboons. Data were collected at the primatology station of the CNRS (France) and a total of 47 bipedal strides were extracted for the present analysis. RESULTS: Adults and adolescents walk bipedally in the same way, and the average kinematic pattern is similar across the age-classes. Infants walk bipedally with longer duty factor, they present larger movement amplitude of the thigh and the amplitude of the knee joint decreases with speed. In contrast, older baboons increase the amplitude of the knee and ankle joints with speed. DISCUSSION: In a non-adapted biped, the postural adjustments of bipedal walking vary with age. In infant baboons, the balance requirements are likely to be higher and these are solved by adopting a "blocking strategy". In older baboons, the postural adjustments are focused on the lower limb and the movements increase with speed. These results may echo, in some respects, the developmental sequence of the intersegmental coordination described in the ontogeny of human locomotion.

Evidence of the use of soft footwear in the Gravettian cave of Cussac (Dordogne, France).

Humans appear to have regularly worn footwear since at least the Early Upper Palaeolithic. However, due to the perishable nature of footwear, the archaeological record of its presence during the Pleistocene is poor. While footwear would have played an essential role in protecting the foot, it could also have been used as ornamentation and/or as a social marker. Footprints may provide the most relevant insight regarding the origin and function of footwear. Here we report the discovery of footprints in Cussac Cave (southwest France) at 28-31 ka cal BP and the results of a multi-focal approach, including experimentation, that demonstrate that Gravettian people most likely wore footwear while moving through the cave. These singular footprints would constitute one of the oldest cases of indirect evidence for this unusual practice in decorated Palaeolithic caves and reinforce the exceptional nature of Cussac already attested by the presence of monumental engravings and funerary deposits.

The quadrupedal walking gait of the olive baboon, Papio anubis: an exploratory study integrating kinematics and EMG.

Primates exhibit unusual quadrupedal features (e.g. diagonal gaits, compliant walk) compared with other quadrupedal mammals. Their origin and diversification in arboreal habitats have certainly shaped the mechanics of their walking pattern to meet the functional requirements necessary for balance control in unstable and discontinuous environments. In turn, the requirements for mechanical stability probably conflict with mechanical energy exchange. In order to investigate these aspects, we conducted an integrative study on quadrupedal walking in the olive baboon (Papio anubis) at the Primatology station of the CNRS in France. Based on kinematics, we describe the centre of mass mechanics of the normal quadrupedal gait performed on the ground, as well as in different gait and substrate contexts. In addition, we studied the muscular activity of six hindlimb muscles using non-invasive surface probes. Our results show that baboons can rely on an inverted pendulum-like exchange of energy (57% on average, with a maximal observed value of 84%) when walking slowly (<0.9 m s-1) with a tight limb phase (∼55%) on the ground using diagonal sequence gaits. In this context, the muscular activity is similar to that of other quadrupedal mammals, thus reflecting the primary functions of the muscles for limb movement and support. In contrast, walking on a suspended branch generates kinematic and muscular adjustments to ensure better control and to maintain stability. Finally, walking using the lateral sequence gait increases muscular effort and reduces the potential for high recovery rates. The present exploratory study thus supports the assumption that primates are able to make use of an inverted pendulum mechanism on the ground using a diagonal walking gait, yet a different footfall pattern and substrate appear to influence muscular effort and efficiency.

Increased performance in juvenile baboons is consistent with ontogenetic changes in morphology.

OBJECTIVES: In many primates, the greater proportion of climbing and suspensory behaviors in the juvenile repertoire likely necessitates good grasping capacities. Here, we tested whether very young individuals show near-maximal levels of grasping strength, and whether such an early onset of grasping performance could be explained by ontogenetic variability in the morphology of the limbs in baboons. MATERIAL AND METHODS: We quantified a performance trait, hand pull strength, at the juvenile and adult stages in a cross-sectional sample of 15 olive baboons (Papio anubis). We also quantified bone dimensions (i.e., lengths, widths, and heights) of the fore- (n = 25) and hind limb (n = 21) elements based on osteological collections covering the whole development of olive baboons. RESULTS: One-year old individuals demonstrated very high pull strengths (i.e., 200% of the adult performance, relative to body mass), that are consistent with relatively wider phalanges and digit joints in juveniles. The mature proportions and shape of the forelimb elements appeared only at full adulthood (i.e., ≥4.5 years), whereas the mature hind limb proportions and shape were observed much earlier during development. DISCUSSION: These changes in limb performance and morphology across ontogeny may be explained with regard to behavioral transitions that olive baboons experience during their development. Our findings highlight the effect of infant clinging to mother, an often-neglected feature when discussing the origins of grasping in primates. The differences in growth patterns, we found between the forelimb and the hind limb further illustrate their different functional roles, having likely evolved under different ecological pressures (manipulation and locomotion, respectively).

The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France).

Footprints represent a unique snapshot of hominin life. They provide information on the size and composition of groups that differs from osteological and archeological remains, whose contemporaneity is difficult to establish. We report here on the discovery of 257 footprints dated to 80,000 y from the Paleolithic site at Le Rozel (Normandy, France), which represent the largest known Neandertal ichnological assemblage to date. We investigate the size and composition of a track-maker group from this large set by developing a morphometric method based on experimental footprints. Our analyses indicate that the footprints were made by a small group comprising different age classes, from early childhood to adult, with a majority of children. The Le Rozel footprints thus provide direct evidence for the size and composition of a Neandertal social group.

The gibbon's Achilles tendon revisited: consequences for the evolution of the great apes?

The well-developed Achilles tendon in humans is generally interpreted as an adaptation for mechanical energy storage and reuse during cyclic locomotion. All other extant great apes have a short tendon and long-fibred triceps surae, which is thought to be beneficial for locomotion in a complex arboreal habitat as this morphology enables a large range of motion. Surprisingly, highly arboreal gibbons show a more human-like triceps surae with a long Achilles tendon. Evidence for a spring-like function similar to humans is not conclusive. We revisit and integrate our anatomical and biomechanical data to calculate the energy that can be recovered from the recoiling Achilles tendon during ankle plantar flexion in bipedal gibbons. Only 7.5% of the required external positive work in a stride can come from tendon recoil, yet it is delivered at an instant when the whole-body energy level drops. Consequently, an additional similar amount of mechanical energy must simultaneously dissipate elsewhere in the system. Altogether, this challenges the concept of an energy-saving function in the gibbon's Achilles tendon. Cercopithecids, sister group of the apes, also have a human-like triceps surae. Therefore, a well-developed Achilles tendon, present in the last common 'Cercopithecoidea-Hominoidea' ancestor, seems plausible. If so, the gibbon's anatomy represents an evolutionary relict (no harm-no benefit), and the large Achilles tendon is not the premised key adaptation in humans (although the spring-like function may have further improved during evolution). Moreover, the triceps surae anatomy of extant non-human great apes must be a convergence, related to muscle control and range of motion. This perspective accords with the suggestions put forward in the literature that the last common hominoid ancestor was not necessarily great ape-like, but might have been more similar to the small-bodied catarrhines.

Behavioral implications of ontogenetic changes in intrinsic hand and foot proportions in olive baboons (Papio Anubis).

OBJECTIVES: Relatively long digits are considered to enhance grasping performance in primates. We tested whether growth-related changes in intrinsic hand and foot proportions may have behavioral implications for growing animals, by examining whether ontogenetic changes in digital proportions are related to variation in voluntary grasping behaviors in baboons. MATERIALS AND METHODS: Longitudinal morphological and behavioral data were collected on 6 captive olive baboons (Papio anubis) as they aged from 5 to 22 months. The length of digits and metapodials, measured from radiographs, were used to calculate phalangeal indices (i.e., PIs: summed length of non-distal phalanges relative to corresponding metapodial length). We also examined the allometric scaling of digital bones relative to body mass. We observed baboon positional behaviors over a 15-day period following the radiographic sessions, quantifying the frequency of forelimb and hindlimb grasping behaviors. RESULTS: PIs for all digits declined during growth, a result of the differential scaling of metapodials (which scaled to body mass with isometry) versus phalanges (which scaled with negative allometry). The incidence of forelimb and hindlimb grasping behaviors declined with age. Though we found no relationship between forelimb grasping and hand proportions, the incidence of hindlimb grasping was directly correlated with postaxial digit PIs. DISCUSSION: Only changes in the intrinsic proportions of the pedal digits are associated with variation in grasping activity in growing baboons. This finding accords previous biomechanical and neuroanatomical studies showing distinct functional roles for the hands and feet during primate locomotion, and has important implications for reconstructing primate locomotor evolution.

The origin of bipedality as the result of a developmental by-product: The case study of the olive baboon (Papio anubis).

In this paper, we point to the importance of considering infancy in the emergence of new locomotor modes during evolution, and particularly when considering bipedal walking. Indeed, because infant primates commonly exhibit a more diverse posturo-locomotor repertoire than adults, the developmental processes of locomotion represent an important source of variation upon which natural selection may act. We have had the opportunity to follow the development of locomotion in captive individuals of a committed quadrupedal primate, the olive baboon (Papio anubis). We observed six infants at two different stages of their development. In total, we were able to analyze the temporal parameters of 65 bipedal steps, as well as their behavioral components. Our results show that while the basic temporal aspects of the bipedal walking gait (i.e., duty factor, dimensionless frequency, and hind lag) do not change during development, the baboon is able to significantly improve the coordination pattern between hind limbs. This probably influences the bout duration of spontaneous bipedal walking. During the same developmental stage, the interlimb coordination in quadrupedal walking is improved and the proportion of quadrupedal behaviors increases significantly. Therefore, the quadrupedal pattern of primates does not impede the developmental acquisition of bipedal behaviors. This may suggest that the same basic mechanism is responsible for controlling bipedal and quadrupedal locomotion, i.e., that in non-human primates, the neural networks for quadrupedal locomotion are also employed to perform (occasional) bipedal walking. In this context, a secondary locomotor mode (e.g., bipedalism) experienced during infancy as a by-product of locomotor development may lead to evolutionary novelties when under appropriate selective pressures.

Segmental morphometrics of the olive baboon (Papio anubis): a longitudinal study from birth to adulthood.

The linear dimensions and inertial characteristics of the body are important in locomotion and they change considerably during the ontogeny of animals, including humans. This longitudinal and ontogenetic study has produced the largest dataset to date of segmental morphometrics in a Catarrhini species, the olive baboon. The objectives of the study were to quantify the changes in body linear and inertial dimensions and to explore their (theoretical) mechanical significance for locomotion. We took full-body measurements of captive individuals at regular intervals. Altogether, 14 females and 16 males were followed over a 7-year period, i.e. from infancy to adulthood. Our results show that individual patterns of growth are very consistent and follow the general growth pattern previously described in olive baboons. Furthermore, we obtained similar growth curve structures for segment lengths and masses, although the respective time scales were slightly different. The most significant changes in body morphometrics occurred during the first 2 years of life and concerned the distal parts of the body. Females and males were similar in size and shape at birth. The rate and duration of growth produced substantial size-related differences throughout ontogeny, while body shapes remained very similar between the sexes. We also observed significant age-related variations in limb composition, with a proximal shift of the centre of mass within the limbs, mainly due to changes in mass distribution and in the length of distal segments. Finally, we observed what we hypothesize to be 'early biomechanical optimization' of the limbs for quadrupedal walking. This is due to a high degree of convergence between the limbs' natural pendular periods in infants, which may facilitate the onset of quadrupedal walking. Furthermore, the mechanical significance of the morphological changes observed in growing baboons may be related to changing functional demands with the onset of autonomous (quadrupedal) locomotion. From a wider perspective, these data provide unique insights into questions surrounding both the processes of locomotor development in primates and how these processes might evolve.

Effect of body mass distribution on the ontogeny of positional behaviors in non-human primates: Longitudinal follow-up of infant captive olive baboons (Papio anubis).

The diversity of primates' positional capabilities is unique among mammals. Indeed, they exhibit a daily repertoire composed of various locomotor and postural modes that may be linked to their particular morphological pattern. Because ontogeny undergoes parallel behavioral and morphological modifications, it may be useful to investigate the biomechanical consequences of the changing body shape. We, therefore, collected accurate quantitative and longitudinal data on positional behaviors, body mass distribution patterns, activities, and environment on a sample of six infant olive baboons, Papio anubis. These baboons are kept at the Primatology Station of the CNRS, France, where they live within the same social group. Individual behaviors were quantified using the focal sampling method. The body mass distribution was estimated according to a geometric model based on direct external measurements. Multivariate analysis enabled us to analyze the interactions between the data. Our results show that body mass distribution changes together with the ontogenetic changes in positional behaviors. At an early age, individuals have distally heavy segment masses in the limbs and an important fraction of the behavioral repertoire involves efficient grasping abilities. At the end of infancy, the same individuals have relatively more mass in proximal segments of the limbs and the proportion of quadrupedal walking is significantly higher while other climbing and suspensory behaviors decreased substantially. The present study experimentally confirms the association between body mass distribution and the positional repertoire of primates. These relationships, when interpreted in the context of basic biomechanical concepts, may improve our understanding of primate locomotion. We discuss further the implications of these functional relationships when modeling the evolutionary pathway of primates. Am. J. Primatol. 78:1201-1221, 2016. © 2016 Wiley Periodicals, Inc.

Bipedality from locomotor autonomy to adulthood in captive olive baboon (Papio anubis): Cross-sectional follow-up and first insight into the impact of body mass distribution.

OBJECTIVE: Despite that the biomechanics of standing and walking bipedally has been extensively studied in nonhuman primates, the morphological features that may constrain, or facilitate, the control of balance and thus of the spontaneous occurrence of bipedal behavior are poorly known. We aim to test the relationship between body mass distribution and bipedal behavior using a nonhuman primate species, the olive baboon, Papio anubis, raised in captivity. MATERIALS AND METHODS: We collected quantitative data on the frequency and duration of bipedalism together with morphometrics on a sample of 22 individuals. We used ontogenetic changes as a natural experiment that provides insights into the impact of morphology. Specifically we focus on 1) quantifying how body mass distribution changes from infancy to adulthood in baboons; and 2) whether the different patterns of mass distribution influence the behavioral variables, i.e., a) the frequency and b) the duration of bouts of bipedal behavior realized in different activity contexts. RESULTS: With regard to assisted bipedal behaviors, the duration and frequency of bouts of standing, contrary to walking, are significantly related to age. With regard to unassisted bipedal behaviors, no correlation to age is observed; the bout duration of walking is strongly correlated to body mass and mass distribution, contrary to the frequency of walking as well as the bout duration and frequency of bipedal standing. DISCUSSION: Our results suggest a close relationship between the pattern of mass distribution and the mechanism of balance control in the spontaneous bipedal walking of baboons. The mechanical effects of the pattern of mass distribution on the ability to perform bipedally in extant nonhuman primates are discussed in the context of the evolution toward habitual bipedalism.

Kinematics and spatiotemporal parameters of infant-carrying in olive baboons.

In the field of biomechanics of quadrupedal locomotion in primates, infant-carrying has received little attention. This study presents the first biomechanical study of infant-carrying in captive female olive baboons (Papio anubis). We test whether females carrying infants conform 1) to the Support Polygon Model (Rollinson and Martin: Symp Zool Soc Lond 48 (1981) 377-427) of gait selection, according to which diagonality should decrease when the infant is carried cranially and increase when the infant is carried dorsally and caudally; 2) to Biewener's (Biewener: Science 245 () 45-48) theory of limb postures, according to which females should extend their hind limbs more due to infant load, especially in the later stages when the infant is not fully autonomous but relatively heavy. This study focuses on the sagittal kinematics of quadrupedal gaits (joint angles and spatiotemporal parameters) of four females with and without infant loads at the CNRS Primatology Station (France). High-speed video recordings were made using the technical platform "Motion Analysis of Primates" available in the animals' place of life. Regarding diagonality, our results do not fully conform to those predicted by the Support Polygon Model of gait selection; however, the model cannot be rejected at this stage in experiment. With regard to limb posture, our results do not support Biewener's (Biewener: Science 245 () 45-48) theory: loaded females do not extend their hind limbs more as predicted; on the contrary, the hind limbs tend to be more flexed when the infant they carry is relatively heavy.

Bipedal behaviour in olive baboons: infants versus adults in a captive environment.

The olive baboon is described as a committed quadrupedal primate. However, available data show that they actually use a variety of locomotor and postural modes. Bipedalism is observed occasionally but spontaneously in captivity and in the wild. As observed in other Catarrhini, immature baboons appear to be more bipedal than adults: this study aims to provide the necessary quantitative data to support this hypothesis, as none has been available so far. The locomotor and postural repertoire was quantified for two age classes: infants beginning to forage independently, and adults. Our results show that infants appear to have a wider repertoire than adults, and bipedal postures and locomotion in infants, although infrequent, appear to distinguish them clearly from adults. In captivity, behavioural context and morphology are the two main factors that could explain age-related positional differences, given a constant ecological context.

Carpal kinematics in quadrupedal monkeys: towards a better understanding of wrist morphology and function.

The purpose of this study is to provide new data on carpal kinematics in primates in order to deepen our understanding of the relationships between wrist morphology and function. To that end, we provide preliminary data on carpal kinematics in seven species of quadrupedal monkeys that have not been previously investigated in this regard (cercopithecoids, n = 4; ceboids, n = 3). We radiographed wrists from cadavers at their maximum radial and ulnar deviations, as well as at maximum flexion and extension. We took angular measurements to quantify the contribution of the mobility of the two main wrist joints (antebrachiocarpal and midcarpal) with respect to total wrist mobility. We also recorded qualitative observations. Our quantitative results show few clear differences among quadrupedal monkeys for radioulnar deviation and flexion-extension: all the primates studied exhibit a greater midcarpal mobility (approximately 54-83% of the total range of motion) than antebrachiocarpal mobility; however, we identified two patterns of carpal kinematics that show the functional impact of previously recognised morphological variations in quadrupedal monkeys. Firstly, qualitative results show that the partition that divides the proximal joint of the wrist in ceboids results in less mobility and more stability of the ulnar part of the wrist than is seen in cercopithecoids. Secondly, we show that the olive baboon specimen (Papio anubis) is characterised by limited antebrachiocarpal mobility for extension; this effect is likely the result of a radial process that projects on the scaphoid notch, as well as an intraarticular meniscus. Because of these close relationships between carpal kinematics and morphology in quadrupedal monkeys, we hypothesise that, to some extent, these functional tendencies are related to their locomotor hand postures.

From bone to plausible bipedal locomotion. Part II: Complete motion synthesis for bipedal primates.

This paper addresses the problem of synthesizing plausible bipedal locomotion according to 3D anatomical reconstruction and general hypotheses on human motion control strategies. In a previous paper [Nicolas, G., Multon, F., Berillon, G., Marchal, F., 2007. From bone to plausible bipedal locomotion using inverse kinematics. Journal of Biomechanics 40 (5) 1048-1057], we have validated a method based on using inverse kinematics to obtain plausible lower-limb motions knowing the trajectory of the ankle. In this paper, we propose a more general approach that also involves computing a plausible trajectory of the ankles for a given skeleton. The inputs are the anatomical descriptions of the bipedal species, imposed footprints and a rest posture. This process is based on optimizing a reference ankle trajectory until a set of criteria is minimized. This optimization loop is based on the assumption that a plausible motion is supposed to have little internal mechanical work and should be as less jerky as possible. For each tested ankle trajectory, inverse kinematics is used to compute a lower-body motion that enables us to compute the resulting mechanical work and jerk. This method was tested on a set of modern humans (male and female, with various anthropometric properties). We show that the results obtained with this method are close to experimental data for most of the subjects. We also demonstrate that the method is not sensitive to the choice of the reference ankle trajectory; any ankle trajectory leads to very similar result. We finally apply the method to a skeleton of Pan paniscus (Bonobo), and compare the resulting motion to those described by zoologists.