Locomotor Repertoire Research Articles

Subterranean to submarine: stress-induced locomotor repertoire expansion and aquatic escape in the Anatolian mole rat (Nannospalax xanthodon) under risk of predation

Subterranean to submarine: stress-induced locomotor repertoire expansion and aquatic escape in the Anatolian mole rat (Nannospalax xanthodon) under risk of predation

Trabecular architecture of the distal femur in extant hominids.

Extant great apes are characterized by a wide range of locomotor, postural and manipulative behaviours that each require the limbs to be used in different ways. In addition to external bone morphology, comparative investigation of trabecular bone, which (re-)models to reflect loads incurred during life, can provide novel insights into bone functional adaptation. Here, we use canonical holistic morphometric analysis (cHMA) to analyse the trabecular morphology in the distal femoral epiphysis of Homo sapiens (n = 26), Gorilla gorilla (n = 14), Pan troglodytes (n = 15) and Pongo sp. (n = 9). We test two predictions: (1) that differing locomotor behaviours will be reflected in differing trabecular architecture of the distal femur across Homo, Pan, Gorilla and Pongo; (2) that trabecular architecture will significantly differ between male and female Gorilla due to their different levels of arboreality but not between male and female Pan or Homo based on previous studies of locomotor behaviours. Results indicate that trabecular architecture differs among extant great apes based on their locomotor repertoires. The relative bone volume and degree of anisotropy patterns found reflect habitual use of extended knee postures during bipedalism in Homo, and habitual use of flexed knee posture during terrestrial and arboreal locomotion in Pan and Gorilla. Trabecular architecture in Pongo is consistent with a highly mobile knee joint that may vary in posture from extension to full flexion. Within Gorilla, trabecular architecture suggests a different loading of knee in extension/flexion between females and males, but no sex differences were found in Pan or Homo, supporting our predictions. Inter- and intra-specific variation in trabecular architecture of distal femur provides a comparative context to interpret knee postures and, in turn, locomotor behaviours in fossil hominins.

Lufengpithecus inner ear provides evidence of a common locomotor repertoire ancestral to human bipedalism

Various lines of evidence have been used to infer the origin of human bipedalism, but the paucity of hominoid postcranial fossils and the diversity of inferred locomotor modes have tended to confound the reconstruction of ancestral morphotypes. Examination of the bony labyrinth morphology of the inner ear of extinct and living hominoids provides independent evidence for inferring the evolution of hominoid locomotor patterns. New computed tomography data and morphometric analyses of the Late Miocene ape Lufengpithecus indicate that it and other stem great apes possess labyrinths similar to one another and show that hominoids initially evolved from a positional repertoire that included orthogrady, below-branch forelimb suspension and progression, above-branch bipedalism, climbing, clambering, and leaping (hylobatid-like) to one that comprised above-branch quadrupedalism, below-branch forelimb suspension, vertical climbing, limited leaping, terrestrial quadrupedal running and walking, possibly with knuckle walking, and short bouts of bipedalism (chimpanzee-like). The bony labyrinth morphology of Lufengpithecus indicates that it probably conforms more closely to the last common ancestors of crown hominoids and hominids in its locomotor behavior than do other Miocene hominoids. Human bipedalism evolved from this common archetypal Lufengpithecus-like locomotor repertoire. The low evolutionary rate of semicircular canal morphology suggests that Lufengpithecus experienced a relative stasis in locomotor behavior, probably due to the uplift of the Tibetan Plateau, which created a stable environment in the Miocene of southwestern China.

New material of Lophiparamys debequensis from the Willwood Formation (early Eocene) of Wyoming, including the first postcrania of the genus

Abstract We report new material of the rare early Eocene rodent Lophiparamys debequensis Wood, 1962 from the Willwood Formation of the southern Bighorn Basin, north-central Wyoming. The new material constitutes the first record of L. debequensis from the Bighorn Basin and documents aspects of the anatomy of Lophiparamys that were previously unknown, including a portion of the maxilla and a portion of the tarsus. The maxillary fragment demonstrates that Lophiparamys has a small P3 and a relatively large infraorbital canal. The tarsus of Lophiparamys is similar to that of other early rodents but differs in a few features that suggest an arboreal locomotor repertoire, including an asymmetric astragalar trochlea, long astragalar neck, transverse astragalar sustentacular facet, short calcaneal tuber, elongate calcaneal ectal facet, and circular calcaneal cuboid facet. The presence of arboreally adapted features in the tarsus of Lophiparamys is consistent with a hypothesized relationship between small-bodied Eocene microparamyine rodents and extant Gliridae. Phylogenetic analysis fails to consistently support this relationship or monophyly of Microparamyinae, but both remain plausible. Comparison of L. debequensis with other species of the genus emphasizes the distinctiveness of L. debequensis and suggests the presence of multiple lineages of Lophiparamys.

Beakiation: how a novel parrot gait expands the locomotor repertoire of living birds.

Occupation of arboreal habitats poses myriad locomotor challenges, driving both anatomical and behavioural innovations across various tetrapod lineages. Here, we report and biomechanically assess a novel, beak-driven locomotor mode-'beakiation'-by which parrots advance along the underside of narrow arboreal substrates. Using high-speed videography and kinetic analyses, we describe the limb loading patterns and pendular mechanics of beakiation, and compare the biomechanical characteristics of this gait with other suspensory behaviours (namely, forelimb-driven brachiation and inverted quadrupedal walking). We report that the parrot beak experiences comparable force magnitudes (approx. 150% body weight in the normal plane; approx. 50% body weight in the fore-aft plane) to the forelimbs of brachiating primates. Parrot beakiation is also characterized by longer-than-expected pendular periods, similar to observations of gibbon brachiation. However, in terms of mechanical energy recovery, beakiation is typified by lower levels of energetic recovery than brachiating specialists: a product of its slower, more careful nature. The observation of this novel behaviour-which adds to a growing base of literature regarding beak-assisted locomotor strategies in birds-highlights the extraordinary behavioural plasticity of birds, the functional versatility of the avian beak, and the difficulties in reconstructing an animal's locomotor repertoire from morphological characteristics alone.

Ulnar morphology in Sahelanthropus tchadensis, what it tells us on the locomotor repertoire of the last common ancestor of humans and

Ulnar morphology in Sahelanthropus tchadensis, what it tells us on the locomotor repertoire of the last common ancestor of humans and

Biomechanical and taxonomic diversity in the Early Pleistocene in East Africa: Structural analysis of a recently discovered femur shaft from Olduvai Gorge (bed I)

Recent Plio-Pleistocene hominin findings have revealed the complexity of human evolutionary history and the difficulties involved in its interpretation. Moreover, the study of hominin long bone remains is particularly problematic, since it commonly depends on the analysis of fragmentary skeletal elements that in many cases are merely represented by small diaphyseal portions and appear in an isolated fashion in the fossil record. Nevertheless, the study of the postcranial skeleton is particularly important to ascertain locomotor patterns. Here we report on the discovery of a robust hominin femoral fragment (OH 84) at the site of Amin Mturi Korongo dated to 1.84 Ma (Olduvai Bed I). External anatomy and internal bone structure of OH 84 were analyzed and compared with previously published data for modern humans and chimpanzees, as well as for Australopithecus, Paranthropus and Homo specimens ranging from the Late Pliocene to Late Pleistocene. Biomechanical analyses based on transverse cross-sections and the comparison of OH 84 with another robust Olduvai specimen (OH 80) suggest that OH 84 might be tentatively allocated to Paranthropus boisei. More importantly, the identification of a unique combination of traits in OH 84 could indicate both terrestrial bipedalism and an arboreal component in the locomotor repertoire of this individual. If interpreted correctly, OH 84 could thus add to the already mounting evidence of substantial locomotor diversity among Early Pleistocene hominins. Likewise, our results also highlight the difficulties in accurately interpreting the link between form and function in the human fossil record based on fragmentary remains, and ultimately in distinguishing between coeval hominin groups due to the heterogeneous pattern of inter- and intraspecific morphological variability detected among fossil femora.

Biomechanics and the origins of human bipedal walking: The last 50 years

Motion analysis, as applied to evolutionary biomechanics, has experienced its own evolution over the last 50 years. Here we review how an ever-increasing fossil record, together with continuing advancements in biomechanics techniques, have shaped our understanding of the origin of upright bipedal walking. The original, and long-established hypothesis held by Lamarck (1809), Darwin (1859) and Keith (1934), amongst others, maintained that bipedality originated in an arboreal context. However, the first field studies of gorilla and chimpanzees from the 1960’s, highlighted their so-called ‘knucklewalking’ quadrupedalism, leading scientists to assume, semi-automatically, that knucklewalking must have been the precursor to bipedality. It would not be until the discovery of skeletons of early human relatives Australopithecus afarensis and Australopithecus prometheus, and the inclusion of methods of analysis from computer science, biomechanics, sports science and medicine, that the knucklewalking hypothesis would be most robustly challenged. Their short, but human-like lower limbs and human-like hand indicated that knucklewalking was not part of our ancestral locomotor repertoire. Rather, most current research in evolutionary biomechanics agrees it was a combination of climbing and bipedalism, both in an arboreal context, which facilitated upright, terrestrial, bipedal walking over short distances.

Early euprimates already had a diverse locomotor repertoire: Evidence from ankle bone morphology

The morphological adaptations of euprimates have been linked to their origin and early evolution in an arboreal environment. However, the ancestral and early locomotor repertoire of this group remains contentious. Although some tarsal bones like the astragalus and the calcaneus have been thoroughly studied, the navicular remains poorly studied despite its potential implications for foot mobility. Here, we evaluate early euprimate locomotion by assessing the shape of the navicular—an important component of the midtarsal region of the foot—using three-dimensional geometric morphometrics in relation to quantified locomotor repertoire in a wide data set of extant primates. We also reconstruct the locomotor repertoire of representatives of the major early primate lineages with a novel phylogenetically informed discriminant analysis and characterize the changes that occurred in the navicular during the archaic primate–euprimate transition. To do so, we included in our study an extensive sample of naviculars (36 specimens) belonging to different species of adapiforms, omomyiforms, and plesiadapiforms. Our results indicate that navicular shape embeds a strong functional signal, allowing us to infer the type of locomotion of extinct primates. We demonstrate that early euprimates displayed a diverse locomotor behavior, although they did not reach the level of specialization of some living forms. Finally, we show that the navicular bone experienced substantial reorganization throughout the archaic primate–euprimate transition, supporting the major functional role of the tarsus during early primate evolution. This study demonstrates that navicular shape can be used as a reliable proxy for primate locomotor behavior. In addition, it sheds light on the diverse locomotor behavior of early primates as well as on the archaic primate–euprimate transition, which involved profound morphological changes within the tarsus, including the navicular bone.

Locomotor adaptations in the Early Miocene species Diamantomys luederitzi (Rodentia, Mammalia) from Uganda (Napak).

The study of morphological adaptations to different ecological parameters among fossil vertebrates has been an important challenge in recent decades. In this paper, we focus on the link between morphological traits and locomotor behavior such as terrestriality, fossoriality and arboreality (including gliding). One of the most diverse groups in which various locomotor habits are represented is rodents, occupying a wide range of ecological niches. This work highlights morphological variations in skulls and humerus in extant rodents with varying locomotion, in order to predict this parameter in the extinct species Diamantomys luederitzi (Early Miocene, Napak, Uganda). Linear Discriminant Analysis and Phylogenetic Flexible Discriminant Analysis are used to analyze datasets obtained via traditional morphometry (measurements) and geometric morphometrics (landmarks). The results show good discrimination between locomotor groups for both structures in extant species: the skull has a wider and longer rostrum in terrestrial and fossorial taxa compared to arboreal rodents, is also higher and posteriorly wider in fossorial taxa; the distal humerus shows elongation of the trochlea and capitulum and a higher trochlea in fossorial and terrestrial species, allowing an increase of stability instead of mobility, which is more important in arboreal taxa for movement in trees. In Diamantomys luederitzi, all skull analyses except one predicted it as a terrestrial species, the other prediction as a glider was possibly linked to the diet. For the distal humerus, this species has been predicted as a terrestrial, fossorial and arboreal taxon in differing analyses, reflected by morphological traits represented in these different locomotor categories. These varying predictions could highlight the intra-specific variation in this fossil species as well as its locomotor repertoire, raising a discussion about the use of different methods in such analyses. In addition to these predictions, several issues are discussed, such as the presence of locomotor signal in the skull and its validity in locomotor studies, as well as the relevance of the use of fragmentary material in such analyses. The results obtained in this work highlight the importance of the locomotor signal in these structures, as well as the possibility of taking into account poorly preserved material, in particular the distal humerus. This article is protected by copyright. All rights reserved.

Three toes and three modes: Dynamics of terrestrial, suspensory, and vertical locomotion in brown-throated three-toed sloths (Bradypodidae, Xenarthra).

Living sloths exhibit numerous anatomical specializations towards inverted quadrupedalism, however, previous studies have noted a more varied locomotor repertoire than previously anticipated. In this study, we present spatiotemporal gait characteristics and triaxial kinetic data from the brown-throated three-toed sloth (Bradypus variegatus) across three locomotor modes: terrestrial quadrupedal "crawling", suspensory walking, and vertical climbing. Compared to quadrupedal crawling and suspensory walking, B. variegatus adopted longer contact times and stride durations, larger duty factors, and greater speed during vertical climbing. Net fore-aft impulses were significantly greater during vertical climbing in both limb pairs than in quadrupedal crawling and suspensory walking. Functionally, during quadrupedal crawling and vertical climbing, both limb pairs served propulsive roles, while differentiation between a propulsive forelimb and braking hindlimb was observed during suspension. Net tangential forces differentiated vertical climbing kinetics from the other modes of locomotion, with the introduction of bidirectional pulling and pushing forces in the forelimb and hindlimb, respectively. The net mediolateral impulses were similar in vertical climbing and quadrupedal crawling as both limb pairs directed forces in one direction, whereas during suspensory walking, the laterally dominant forelimb was opposed by the medially dominant hindlimb. In total, this study provides novel data on the diverse locomotor dynamics in a slow-moving arboreal tetrapod and posits new testable hypotheses about the neuroplasticity and ease of transitioning between locomotor behaviors. The strikingly similar kinetic profiles of quadrupedal crawling and suspensory walking compared to vertical climbing suggest shared neuromuscular and mechanical demands between these mirrored locomotor modes.

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 Jkg-1m-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.

Trabecular bone ontogeny tracks neural development and life history among humans and non-human primates

Trabecular bone-the spongy bone inside marrow cavities-adapts to its mechanical environment during growth and development. Trabecular structure can therefore be interpreted as a functional record of locomotor behavior in extinct vertebrates. In this paper, we expand upon traditional links between form and function by situating ontogenetic trajectories of trabecular bone in four primate species into the broader developmental context of neural development, locomotor control, and ultimately life history. Our aim is to show that trabecular bone structure provides insights into ontogenetic variation in locomotor loading conditions as the product of interactions between increases in body mass and neuromuscular maturation.Our results demonstrate that age-related changes in trabecular bone volume fraction (BV/TV) are strongly and linearly associated with ontogenetic changes in locomotor kinetics. Age-related variation in locomotor kinetics and BV/TV is in turn strongly associated with brain and body size growth in all species. These results imply that age-related variation in BV/TV is a strong proxy for both locomotor kinetics and neuromuscular maturation. Finally, we show that distinct changes in the slope of age-related variation in bone volume fraction correspond to the age of the onset of locomotion and the age of locomotor maturity.Our findings compliment previous studies linking bone development to locomotor mechanics by providing a fundamental link to brain development and life history. This implies that trabecular structure of fossil subadults can be a proxy for the rate of neuromuscular maturation and major life history events like locomotor onset and the achievement of adult-like locomotor repertoires.

Do femoral biomechanical properties follow locomotor changes in primates? An ontogenetic study of olive baboons (Papio anubis)

AbstractObjectivesThe locomotor repertoire of the olive baboon (Papio anubis) changes during ontogeny from a diversified pattern in infancy to a stereotyped pattern in adults. On the assumption that the femoral diaphysis adapts to variations in the loading regime, we aimed to evaluate the impact of locomotor changes on femoral biomechanical properties during the ontogeny of olive baboons.Materials and MethodsWe used the cross‐sectional geometry of 50 three‐dimensional‐reconstructed olive baboon femora of an ontogenetic sample including individuals from birth to 28 years of age and belonging to a single social group. Cross section parameters were extracted, and cortical thickness maps were produced from 20% to 80% of the femoral length.ResultsNewborns present a distribution of cortical thickness that is longitudinally symmetric and anteroposterior shaped cross sections at midshaft. From this condition, infants, juveniles and adults femoral diaphyses are all characterized by a proximal shift in cortical thickness and a mediolateralization of bending rigidity and strength at midshaft.DiscussionAs soon as locomotor autonomy is established and despite a combination of several significant locomotor modes, the biomechanical pattern remains stable during ontogeny. Our results do not reflect the changes in locomotor repertoires that occur during ontogeny of Papio anubis. We therefore advise caution when inferring complex locomotor behavior from femoral biomechanical properties.

De novo establishment of circuit modules restores locomotion after spinal cord injury in adult zebrafish

Mechanisms underlying spontaneous locomotor recovery after spinal cord injury (SCI) remain unclear. Using adult zebrafish with complete SCI, we show that V2a interneurons regrow their axon to bridge the lesioned spinal segments in a subclass-specific and chronological order. Early after SCI, reestablishment of a unitary high-rhythm locomotor circuit is driven merely by axon-regrown fast V2a interneurons. Later, the reestablished intraspinal de novo circuit is organized into a modular design by axon-regrown fast and slow V2a interneurons rostral to the lesion, selectively driving caudal fast V2a/motor neurons and slow V2a/motor neurons, respectively. This orderly circuitry reestablishment determines the stepwise restoration of locomotor repertoire and recapitulates developmental processes. This progress can be interrupted by ablation of calretinin, a fast module-related protein, and accelerated by physical training. These findings suggest that promotion of axon regrowth of propriospinal V2a interneurons and establishment of de novo intraspinal circuits underpin the effectiveness of physical training in patients after SCI.

Postcranial evidence of late Miocene hominin bipedalism in Chad.

Bipedal locomotion is one of the key adaptations that define the hominin clade. Evidence of bipedalism is known from postcranial remains of late Miocene hominins as early as 6 million years ago (Ma) in eastern Africa1-4. Bipedality of Sahelanthropus tchadensis was hitherto inferred about 7 Ma in central Africa (Chad) based on cranial evidence5-7. Here we present postcranial evidence of the locomotor behaviour of S. tchadensis, with new insights into bipedalism at the early stage of hominin evolutionary history. The original material was discovered at locality TM 266 of the Toros-Ménalla fossiliferous area and consists of one left femur and two, right and left, ulnae. The morphology of the femur is most parsimonious with habitual bipedality, and the ulnae preserve evidence of substantial arboreal behaviour. Taken together, these findings suggest that hominins were already bipeds at around 7 Ma but also suggest that arboreal clambering was probably a significant part of their locomotor repertoire.

Pathways to primate hip function.

Understanding how diverse locomotor repertoires evolved in anthropoid primates is key to reconstructing the clade's evolution. Locomotor behaviour is often inferred from proximal femur morphology, yet the relationship of femoral variation to locomotor diversity is poorly understood. Extant acrobatic primates have greater ranges of hip joint mobility—particularly abduction—than those using more stereotyped locomotion, but how bony morphologies of the femur and pelvis interact to produce different locomotor abilities is unknown. We conducted hypothesis-driven path analyses via regularized structural equation modelling (SEM) to determine which morphological traits are the strongest predictors of hip abduction in anthropoid primates. Seven femoral morphological traits and two hip abduction measures were obtained from 25 primate species, split into broad locomotor and taxonomic groups. Through variable selection and fit testing techniques, insignificant predictors were removed to create the most parsimonious final models. Some morphological predictors, such as femur shaft length and neck-shaft angle, were important across models. Different trait combinations best predicted hip abduction by locomotor or taxonomic group, demonstrating group-specific linkages among morphology, mobility and behaviour. Our study illustrates the strength of SEM for identifying biologically important relationships between morphology and performance, which will have future applications for palaeobiological and biomechanical studies.

Femoral neck anteversion in Xenarthra

Degree of femoral neck anteversion distinguishes extinct sloth genera from each other, particularly when analyzed in combination with femoral neck angle. These data have been previously analyzed using a comparative sample including only extant sloths, which limits functional interpretations. Here, we analyze both femoral neck anteversion and femoral neck angle broadly across Xenarthrans to reconstruct the locomotor behavior of extinct Caribbean sloths.Femora of four extinct sloth genera (n=49; Acratocnus; Megalocnus; Neocnus; Parocnus) and nine extant Xenarthra genera (n=63; Bradypus; Choloepus; Cyclopes; Dasypus; Euphractus; Myrmecophaga; Pridontes; Tamandua; Tolypeutes) were surface scanned. Morphological axes representing the femoral neck, shaft, geometric axis of the femoral condyles (GA), and maximum dimension of the femoral shaft cross section at 30% of whole bone length were extracted. Degree of femoral neck anteversion was calculated relative to both the GA and cross section axes. Femoral neck angle was additionally calculated as the angle between the femoral neck and shaft axes. Non‐parametric MANOVAs with post hoc multiple comparison tests were used to test how the genera differed when the angle of femoral neck anteversion and femoral neck angle were analyzed in conjunction.Parocnusdoes not significantly differ from any of the armadillo taxa when femoral neck angle is analyzed with either of the two anteversion angles (p&gt;0.171 for all pairwise comparisons), supporting previous suggestions that Parocnuswas largely terrestrial. This finding also indicates that Parocnus may have had similar hindlimb biomechanics to armadillos, and possibly utilized digging behaviors. Megalocnusand Acratocnusdiffer significantly from all extant Xenarthrans (p&lt;0.043 for all pairwise comparisons), but do not exhibit significantly different morphologies from each other (p=0.911). This suggests that the two genera may have utilized similar hip mechanics; however, their locomotor repertoires cannot be accommodated within the range of those exhibited by extant Xenarthrans.

A new early hominin calcaneus from Kromdraai (South Africa).

The Kromdraai site in South Africa has yielded numerous early hominin fossils since 1938. As a part of recent excavations within Unit P, a largely complete early hominin calcaneus (KW 6302) was discovered. Due to its role in locomotion, the calcaneus has the potential to reveal important form/function relationships. Here, we describe KW 6302 and analyze its preserved morphology relative to human and nonhuman ape calcanei, as well as calcanei attributed to Australopithecus afarensis, Australopithecus africanus, Australopithecus sediba, Homo naledi, and the Omo calcaneus (either Paranthropus or early Homo). KW 6302 calcaneal morphology is assessed using numerous quantitative metrics including linear measures, calcaneal robusticity index, relative lateral plantar process position, Achilles tendon length reconstruction, and a three-dimensional geometric morphometric sliding semilandmark analysis. KW 6302 exhibits an overall calcaneal morphology that is intermediate between humans and nonhuman apes, although closer to modern humans. KW 6302 possesses many traits that indicate it was likely well-adapted for terrestrial bipedal locomotion, including a relatively flat posterior talar facet and a large lateral plantar process that is similarly positioned to modern humans. It also retains traits that indicate that climbing may have remained a part of its locomotor repertoire, such as a relatively gracile tuber and a large peroneal trochlea. Specimens from Kromdraai have been attributed to either Paranthropus robustus or early Homo; however, there are no definitively attributed calcanei for either genus, making it difficult to taxonomically assign this specimen. KW 6302 and the Omo calcaneus, however, fall outside the range of expected variation for an extant genus, indicating that if the Omo calcaneus was Paranthropus, then KW 6302 would likely be attributed to early Homo (or vice versa).

Growth and development of trabecular structure in the calcaneus of Japanese macaques (Macaca fuscata) reflects locomotor behavior, life history, and neuromuscular development.

Bone structure dynamically adapts to its mechanical environment throughout ontogeny by altering the structure of trabecular bone, the three‐dimensional mesh‐like structure found underneath joint surfaces. Trabecular structure, then, can provide a record of variation in loading directions and magnitude; and in ontogenetic samples, it can potentially be used to track developmental shifts in limb posture. We aim to broaden the analysis of trabecular bone ontogeny by incorporating interactions between ontogenetic variation in locomotor repertoire, neuromuscular maturation, and life history. We examine the associations between these variables and age‐related variation in trabecular structure in the calcaneus of Japanese macaques (Macaca fuscata). We used high‐resolution micro‐computed tomography scanning to image the calcaneus in a cross‐sectional sample of 34 juvenile M. fuscata aged between 0 and 7 years old at the Primate Research Institute, Japan. We calculated whole bone averages of standard trabecular properties and generated whole‐bone morphometric maps of bone volume fraction and Young’s modulus. Trabecular structure becomes increasingly heterogeneous in older individuals. Bone volume fraction (BV/total volume [TV]) decreases during the first month of life and increases afterward, coinciding with the onset of independent locomotion in M. fuscata. At birth, primary Young’s modulus is oriented orthogonal to the ossification center, but after locomotor onset bone structure becomes stiffest in the direction of joint surfaces and muscle attachments. Age‐related variation in bone volume fraction is best predicted by an interaction between the estimated percentage of adult brain size, body mass, and locomotor onset. To explain our findings, we propose a model where interactions between age‐related increases in body weight and maturation of the neuromuscular system alter the loading environment of the calcaneus, to which the internal trabecular structure dynamically adapts. This model cannot be directly tested based on our cross‐sectional data. However, confirmation of the model by longitudinal experiments and in multiple species would show that trabecular structure can be used both to infer behavior from fossil morphology and serve as a valuable proxy for neuromuscular maturation and life history events like locomotor onset and the achievement of an adult‐like gait. This approach could significantly expand our knowledge of the biology and behavior of fossil species.