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The morphology of the proximal carpals (scaphoid, lunate, triquetrum) are linked to the range of motion (ROM) at the radiocarpal and midcarpal joints. While the relationship between ROM and habitual locomotor mode is well established, it has yet to be investigated whether relative patterns of internal bone architecture reflect the kinematics and kinetics at the proximal row. As internal bone is known to model its structure to habitually incurred forces, internal architecture has the potential to provide insight into how a joint(s) have been loaded during the lifetime of an individual. Using a broad sample of extant great apes and humans (n=177 total), this study investigates whether relative differences in the bone volume to total volume (BV/TV) and degree of anisotropy (DA) across the scaphoid, lunate and triquetrum correlate with the presumed force transfer and biomechanics of the hominoid wrist. Results reveal broad patterns in BV/TV and DA differentiated hominoids by their predominant locomotor mode. The human pattern suggests the lunate may be the most highly strained bone within the proximal row. Both knuckle-walking taxa (Gorilla, Pan) exhibited similar architectural patterns suggesting they are adapted to resist similar forces in this region of the wrist. The relatively high DA across all Pongo carpals suggests it may have more stereotypical wrist loading than commonly assumed. Finally, the distinctly low DA in the triquetrum across all taxa suggests force transfer via the synapomorphic Triangular Fibrocartilage Complex may leave a distinctive signature in the internal bone architecture that requires further investigation.

Bipedalism, a defining feature of the human lineage, is thought to have evolved as forests retreated in the late Miocene-Pliocene. Chimpanzees living in analogous habitats to early hominins offer a unique opportunity to investigate the ecological drivers of bipedalism that cannot be addressed via the fossil record alone. We investigated positional behavior and terrestriality in a savanna-mosaic community of chimpanzees ( Pan troglodytes schweinfurthii ) in the Issa Valley, Tanzania as the first test in a living ape of the hypothesis that wooded, savanna habitats were a catalyst for terrestrial bipedalism. Contrary to widely accepted hypotheses of increased terrestriality selecting for habitual bipedalism, results indicate that trees remained an essential component of the hominin adaptive niche, with bipedalism evolving in an arboreal context, likely driven by foraging strategy.

In 1994, Hunt published the 'postural feeding hypothesis'-a seminal paper on the origins of hominin bipedalism-founded on the detailed study of chimpanzee positional behavior and the functional inferences derived from the upper and lower limb morphology of the Australopithecus afarensis A.L. 288-1 partial skeleton. Hunt proposed a model for understanding the potential selective pressures on hominins, made robust, testable predictions based on Au. afarensis functional morphology, and presented a hypothesis that aimed to explain the dual functional signals of the Au. afarensis and, more generally, early hominin postcranium. Here we synthesize what we have learned about Au. afarensis functional morphology and the dual functional signals of two new australopith discoveries with relatively complete skeletons (Australopithecus sediba and StW 573 'Australopithecus prometheus'). We follow this with a discussion of three research approaches that have been developed for the purpose of drawing behavioral inferences in early hominins: (1) developments in the study of extant apes as models for understanding hominin origins; (2) novel and continued developments to quantify bipedal gait and locomotor economy in extant primates to infer the locomotor costs from the anatomy of fossil taxa; and (3) novel developments in the study of internal bone structure to extract functional signals from fossil remains. In conclusion of this review, we discuss some of the inherent challenges of the approaches and methodologies adopted to reconstruct the locomotor modes and behavioral repertoires in extinct primate taxa, and notably the assessment of habitual terrestrial bipedalism in early hominins.

La main humaine a changé de forme et de fonction tout au long de notre évolution en raison, entre autres, de la bipédie terrestre, de l’utilisation d’outils et de l’asymétrie directionnelle telle que la préférence manuelle. Cependant, nous manquons d’informations sur la variabilité potentielle des facteurs clés pouvant affecter la force de préhension et la dextérité, en particulier la taille, la forme et la fonction quotidienne de la main. Nous avons mesuré la force de préhension par pincemen...

The trabecular bone morphology of adult extant primates has been shown to reflect mechanical loading related to locomotion. However, ontogenetic studies of humans and other mammals suggest an adaptive lag between trabecular bone response and current mechanical loading patterns that could result in adult trabecular bone mor-phology reflecting juvenile behaviours. This study investigates ontogenetic changes in the trabecular bone structure of the third metacarpal of mountain gorillas (Gorilla beringei beringei; n= 26) and western lowland gorillas (Gorilla gorilla gorilla; n= 26) and its relationship to expected changes in locomotor loading patterns. Results show that trabecular bone reflects predicted mechanical loading throughout ontogeny. Bone volume fraction, trabecular thickness and trabecular number are low at birth and in-crease with age, although degree of anisotropy remains relatively stable throughout ontogeny. A high concentration of bone volume fraction can be observed in the dis-topalmar region of the third metacarpal epiphysis in early ontogeny, consistent with the high frequency of climbing, suspensory and other grasping behaviours in young gorillas. High trabecular bone concentration increases dorsally in the epiphysis dur-ing the juvenile period as terrestrial knuckle-walking becomes the primary form of locomotion. However, fusion of the epiphysis does not take place until 10–11 years of age, and overall trabecular structure does not fully reflect the adult pattern until 12 years of age, indicating a lag between adult- like behaviours and adult- like tra -becular morphology. We found minimal differences in trabecular ontogeny between mountain and western lowland gorillas, despite presumed variation in the frequencies of arboreal locomotor behaviours. Altogether, ontogenetic changes in Gorilla metacar-pal trabecular structure reflect overall genus- level changes in locomotor behaviours throughout development, but with some ontogenetic lag that should be considered when drawing functional conclusions from bone structure in extant or fossil adoles-cent specimens

Summary Modelling a muscle key to flexing the thumb, a new study suggests that the powerful opposability that characterises the dextrous human hand evolved in some of our fossil relatives about two million years ago — a time when tool use became more important.

Although hand grip strength is critical to the daily lives of humans and our arboreal great ape relatives, the human hand has changed in form and function throughout our evolution due to terrestrial bipedalism, tool use, and directional asymmetry (DA) such as handedness. Here we investigate how hand form and function interact in modern humans to gain an insight into our evolutionary past. We measured grip strength in a heterogeneous, cross-sectional sample of human participants (n = 662, 17 to 83 years old) to test the potential effects of age, sex, asymmetry (hand dominance and handedness), hand shape, occupation, and practice of sports and musical instruments that involve the hand(s). We found a significant effect of sex and hand dominance on grip strength, but not of handedness, while hand shape and age had a greater influence on female grip strength. Females were significantly weaker with age, but grip strength in females with large hands was less affected than those with long hands. Frequent engagement in hand sports significantly increased grip strength in the non-dominant hand in both sexes, while only males showed a significant effect of occupation, indicating different patterns of hand dominance asymmetries and hand function. These results improve our understanding of the link between form and function in both hands and offer an insight into the evolution of human laterality and dexterity.

Understanding the manual abilities of fossil hominins has been a focus of palaeoanthropological research for decades. Of interest are the morphological characteristics of the thumb due to its fundamental role in manipulation, particularly that of the trapeziometacarpal joint. Considerably less attention has been given to the thumb metacarpophalangeal (MCP) joint, which plays a role in stabilizing the thumb during forceful grasps and precision pinching. In this study we use a three-dimensional geometric morphometric approach to quantify the shape of the first metacarpal head in extant hominids (Homo, Pan, Gorilla and Pongo) and six fossil hominin species (Homo neanderthalensis Tabun C1 and La Chappelle-aux-Saints, Homo naledi U.W. 101-1282, Australopithecus sediba MH2, Paranthropus robustus/early Homo SK84, Australopithecus africanus StW 418, Australopithecus afarensis A.L. 333w-39), with the aims of identifying shapes that may be correlated with human-like forceful opposition and determining if similar morphologies are present in fossil hominins. Results show that humans differ from extant great apes by having a distally flatter articular surface, larger epicondyle surface area, and a larger radial palmar condyle. We suggest that this suite of features is correlated with a lower range of motion at the MCP joint, which would enhance the thumbs ability to resist the elevated loads associated with the forceful precision grips typical of humans. Great ape genera are each differentiated by distinctive morphological features, each of which is consistently correlated with the predicted biomechanical demands of their particular locomotor and/or manipulatory habits. Neanderthals and U.W. 101-1282 fall within the modern human range of variation, StW 418, SK 84 and U.W. 88-119 fall in between humans and great apes, and A.L. 333w-39 falls within Pan variation. These results agree with those of traditional linear analyses while providing a more comprehensive quantitative basis from which to interpret the hand functional morphology of extinct hominins.

The suite of anatomical features contributing to the unique gripping capabilities of the modern human hand evolved alongside the proliferation of Lower Palaeolithic flaked tool technologies across the Old World. Experimental studies investigating their potential co-evolution suggest that the use of flakes, handaxes, and other stone tools is facilitated by manipulative capabilities consistent with the evolutionary trajectory of the hominin hand during this period. Grip analyses have provided important contributions to this understanding. To date, however, there has been no large-scale investigation of grip diversity during flaked stone-tool use, empirical comparative analyses of grip use frequencies, or examination of ergonomic relationships between grip choice and stone tool type and form.\ud \ud Here, we conduct four experimental studies, using replica Lower Palaeolithic stone tools in a series of actualistic and laboratory-based contexts, to record grip type and frequency of grip use during 1067 stone tool-use events by 123 individuals. Using detailed morphometric data recorded from each tool, we demonstrate how grip choice varies according to the type and form of stone tool used, and how these relationships differ between tool-use contexts. We identify 29 grip types across all tool-use events, with significant differences recorded in their frequency of use dependent on tool type, tool form, and the context of use. Despite the influence of these three factors, there is consistency in the frequent use of a limited number (?4) of grip types within each experiment and the consistent and seemingly forceful recruitment of the thumb and index finger. Accordingly, we argue that there are deep-rooted, ergonomically-related, regularities in how stone tools are gripped during their use, that these regularities may have been present during the use of stone tools by Plio-Pleistocene hominins, and any subsequent selective pressures would likely have been focused on the first and second digit.

Homo naledi fossils from the Rising Star cave system provide important insights into the diversity of hand morphology within the genus Homo. Notably, the pollical (thumb) metacarpal (Mc1) displays an unusual suite of characteristics including a median longitudinal crest, a narrow proximal base, and broad flaring intrinsic muscle flanges. The present study evaluates the affinities of H. naledi Mc1 morphology via 3D geometric morphometric analysis of shaft shape using a broader comparative sample (n = 337) of fossil hominins, recent humans, apes, and cercopithecoid monkeys than in prior work. Results confirm that the H. naledi Mc1 is distinctive from most other hominins in being narrow at the proximal end but surmounted by flaring muscle flanges distally. Only StW 418 (Australopithecus cf. africanus) is similar in these aspects of shape. The gracile proximal shaft is most similar to cercopithecoids, Pan, Pongo, Australopithecus afarensis, and Australopithecus sediba, suggesting that H. naledi retains the condition primitive for the genus Homo. In contrast, Neandertal Mc1s are characterized by wide proximal bases and shafts, pinched midshafts, and broad distal flanges, while those of recent humans generally have straight shafts, less robust muscle flanges, and wide proximal shafts/bases. Although uncertainties remain regarding character polarity, the morphology of the H. naledi thumb might be interpreted as a retained intermediate state in a transformation series between the overall gracility of the shaft and the robust shafts of later hominins. Such a model suggests that the addition of broad medial and lateral muscle flanges to a primitively slender shaft was the first modification in transforming the Mc1 into the overall more robust structure exhibited by other Homo taxa including Neandertals and recent Homo sapiens in whose shared lineage the bases and proximal shafts became expanded, possibly as an adaptation to the repeated recruitment of powerful intrinsic pollical muscles.

Haeusler et al. (1) suggest that our analysis (2) of the distribution of relative bone volume across the articular surface (figure 5) does not justify different taxonomic allocations or locomotor classifications. We agree with their first suggestion, and we did not use these data to make direct arguments for the taxonomic attribution of either specimen in our paper. With regard to locomotor classifications, we note that our results “further [support] the inferred differences in loading between these two specimens as evidenced by their internal BV/TV distribution” (2). The strongest evidence for locomotor differences between StW 522 and StW 311 comes from the internal distribution of trabecular bone, which is not captured by the geometric … [↵][1]1To whom correspondence may be addressed. Email: m.skinner{at}kent.ac.uk. [1]: #xref-corresp-1-1

This chapter presents description and analysis of the metacarpals and manual phalanges from Sterkfontein. Although the morphology is generally similar across the sample where there are duplicates of the same element, there are differences in size that are quite remarkable within the context of all South African hominins. Some very large specimens suggest the presence of individuals at Sterkfontein with much larger hands, and presumably larger body size, at Sterkfontein than those of A. sediba MH2, H. naledi and the Swartkrans hominins. Australopithecus africanus had human-like proportions, but this may be plesiomorphic within the hominoid clade. The potentially less mobile trapezium-Mc1 joint, absence of a fully developed palmar pulp on the distal thumb, more limited pronation of the index finger, and potentially more wedge-shaped trapezoid inferred from the preserved external morphology, is consistent with lower manipulative loading of the thumb than is typical of later Homo. As for other forelimb elements, moderately curved manual phalanges suggests a greater reliance on forelimb-dominated locomotor behaviors and perhaps selection for more frequent use of an arboreal environment in A. africanus than is found in A. afarensis. Thus, within this broader context, the Sterkfontein fossil hominin remains are not unusual. The Sterkfontein hand fossils suggest an overall manipulative and locomotor loading regime that was more similar to that of other South African australopiths and distinct from that of later Homo, but more refined functional interpretations require additional fossil evidence, particularly from associated hand skeletons

The analysis of internal trabecular and cortical bone has been an informative tool for drawing inferences about behaviour in extant and fossil primate taxa. Within the hand, metacarpal bone architecture has been shown to correlate well with primate locomotion; however, the extent of morphological differences across taxa is unexpectedly small given the variability in hand use. One explanation for this observation is that the activity-related differences in the joint loads acting on the bone are simply smaller than estimated based on commonly used proxies (i.e. external loading and joint posture), which neglect the influence of muscle forces. In this study, experimental data and a musculoskeletal finger model are used to test this hypothesis by comparing differences between climbing and knuckle-walking locomotion of captive bonobos ( Pan paniscus ) based on (i) joint load magnitude and direction predicted by the models and (ii) proxy estimations. The results showed that the activity-related differences in predicted joint loads are indeed much smaller than the proxies would suggest, with joint load magnitudes being almost identical between the two locomotor modes. Differences in joint load directions were smaller but still evident, indicating that joint load directions might be a more robust indicator of variation in hand use than joint load magnitudes. Overall, this study emphasizes the importance of including muscular forces in the interpretation of skeletal remains and promotes the use of musculoskeletal models for correct functional interpretations.

Objectives: Several studies have investigated potential functional signals in the trabecular structure of the primate proximal humerus but with varied success. Here we apply for the first time a ìwhole-epiphysesî approach to analysing trabecular bone in the humeral head with the aim of providing a more holistic interpretation of trabecular variation in relation to habitual locomotor or manipulative behaviors in several extant primates and Australopithecus africanus. \ud \ud Materials and Methods: We use a ìwhole-epiphysisî methodology in comparison to the traditional volume of interest (VOI) approach to investigate variation in trabecular structure and joint loading in the proximal humerus of extant hominoids, Ateles and A. africanus (StW 328).\ud \ud Results: There are important differences in the quantification of trabecular parameters using a ìwhole-epiphysisî versus a VOI-based approach. Variation in trabecular structure across knucklewalking African apes, suspensory taxa, and modern humans was generally consistent with predictions of load magnitude and inferred joint posture during habitual behaviors. Higher relative trabecular bone volume and more isotropic trabeculae in StW 328 suggest A. africanus may have still used its forelimbs for arboreal locomotion.\ud \ud Discussion: A whole-epiphysis approach to analysing trabecular structure of the proximal humerus can help distinguish functional signals of joint loading across extant primates and can provide novel insight into habitual behaviors of fossil hominins.

Although gorillas rarely use tools in the wild, their manipulative skills during plant processing may be similar to those of other tool-using great apes. Virunga mountain gorillas are known for the complexity in their methods of thistle and nettle plant preparation in the wild. However, there has been no comparable data on food processing in the population of mountain gorillas from the Bwindi Impenetrable National Park, Uganda. We investigated the manual actions and hand grips used when accessing edible parts of two hard-to-process plants defended by stinging hairs, epidermis or periderm (i.e., peel of Urera hypselodendron and pith of Mimulopsis arborescens) and one undefended plant (i.e., leaves of Momordica foetida) in 11 Bwindi wild mountain gorillas (Gorilla beringei beringei) using video records ad libitum. Similar to thistle feeding by Virunga gorillas, Bwindi gorillas used the greatest number of manual actions for the most hard-to-process plant (U. hypselodendron), the actions were ordered in several key stages and organised hierarchically. The demands of processing plant material elicited 19 different grips and variable thumb postures, of which three grips were new and 16 grips have either been previously reported or show clear similarities to grips used by other wild and captive African apes and humans. Moreover, our study only partly supports a functional link between diet and hand morphology in mountain gorillas and suggests that the gorilla hand is best adapted to forceful grasping that is required for both manipulation and arboreal locomotion.

Bone form reflects both the genetic profile and behavioural history of an individual. As cortical bone is able to remodel in response to mechanical stimuli, interspecific differences in cortical bone thickness may relate to loading during locomotion or manual behaviours during object manipulation. Here, we test the application of a novel method of cortical bone mapping to the third metacarpal (Mc3) and talus of Pan, Pongo, and Homo. This method of analysis allows measurement of cortical thickness throughout the bone, and as such is applicable to elements with complex morphology. In addition, it allows for registration of each specimen to a canonical surface, and identifies regions where cortical thickness differs significantly between groups. Cortical bone mapping has potential for application to palaeoanthropological studies; however, due to the complexity of correctly registering homologous regions across varied morphology, further methodological development would be advantageous.

This research was supported by The Max Planck Society and the European Research Council Starting Grant #336301.

Approximately 11.6-million-year-old fossils reveal an ape with arms suited to hanging in trees but human-like legs, suggesting a form of locomotion that might push back the timeline for when walking on two feet evolved. Ancient ape fossil reveals a previously unknown form of locomotion.

Introduction Knowledge of internal finger loading during human and non-human primate activities such as tool use or knuckle-walking has become increasingly important to reconstruct the behaviour of fossil hominins based on bone morphology. Musculoskeletal models have proven useful for predicting these internal loads during human activities, but load predictions for non-human primate activities are missing due to a lack of suitable finger models. The main goal of this study was to implement both a human and a representative non-human primate finger model to facilitate comparative studies on metacarpal bone loading. To ensure that the model predictions are sufficiently accurate, the specific goals were: (1) to identify species-specific model parameters based on in vitro measured fingertip forces resulting from single tendon loading and (2) to evaluate the model accuracy of predicted fingertip forces and net metacarpal bone loading in a different loading scenario. Materials & Methods Three human and one bonobo (Pan paniscus) fingers were tested in vitro using a previously developed experimental setup. The cadaveric fingers were positioned in four static postures and load was applied by attaching weights to the tendons of the finger muscles. For parameter identification, fingertip forces were measured by loading each tendon individually in each posture. For the evaluation of model accuracy, the extrinsic flexor muscles were loaded simultaneously and both the fingertip force and net metacarpal bone force were measured. The finger models were implemented using custom Python scripts. Initial parameters were taken from literature for the human model and own dissection data for the bonobo model. Optimized model parameters were identified by minimizing the error between predicted and experimentally measured fingertip forces. Fingertip forces and net metacarpal bone loading in the combined loading scenario were predicted using the optimized models and the remaining error with respect to the experimental data was evaluated. Results The parameter identification procedure led to minor model adjustments but considerably reduced the error in the predicted fingertip forces (root mean square error reduced from 0.53/0.69 N to 0.11/0.20 N for the human/bonobo model). Both models remained physiologically plausible after the parameter identification. In the combined loading scenario, fingertip and net metacarpal forces were predicted with average directional errors below 6° and magnitude errors below 12%. Conclusions This study presents the first attempt to implement both a human and non-human primate finger model for comparative palaeoanthropological studies. The good agreement between predicted and experimental forces involving the action of extrinsic flexors—which are most relevant for forceful grasping—shows that the models are likely sufficiently accurate for comparisons of internal loads occurring during human and non-human primate manual activities.

Studies of femoral trabecular structure have shown that the orientation and volume of bone are associated with variation in loading and could be informative about individual joint positioning during locomotion. In this study, we analyse for the first time trabecular bone patterns throughout the femoral head using a whole?epiphysis approach to investigate how potential trabecular variation in humans and great apes relates to differences in locomotor modes. Trabecular architecture was analysed using microCT scans of Pan troglodytes (n = 20), Gorilla gorilla (n = 14), Pongo sp. (n = 5) and Homo sapiens (n = 12) in medtool 4.1. Our results revealed differences in bone volume fraction (BV/TV) distribution patterns, as well as overall trabecular parameters of the femoral head between great apes and humans. Pan and Gorilla showed two regions of high BV/TV in the femoral head, consistent with hip posture and loading during two discrete locomotor modes: knuckle?walking and climbing. Most Pongo specimens also displayed two regions of high BV/TV, but these regions were less discrete and there was more variability across the sample. In contrast, Homo showed only one main region of high BV/TV in the femoral head and had the lowest BV/TV, as well as the most anisotropic trabeculae. The Homo trabecular structure is consistent with stereotypical loading with a more extended hip compared with great apes, which is characteristic of modern human bipedalism. Our results suggest that holistic evaluations of femoral head trabecular architecture can reveal previously undetected patterns linked to locomotor behaviour in extant apes and can provide further insight into hip joint loading in fossil hominins and other primates.

Trabecular bone remodels during life in response to loading and thus should, at least in part, reflect potential variation in the magnitude, frequency and direction of joint loading across different hominid species. Here we analyse the trabecular structure across all non-pollical metacarpal distal heads (Mc2-5) in extant great apes, expanding on previous volume of interest and whole-epiphysis analyses that have largely focused on only the first or third metacarpal. Specifically, we employ both a univariate statistical mapping and a multivariate approach to test for both inter-ray and interspecific differences in relative trabecular bone volume fraction (RBV/TV) and degree of anisotropy (DA) in Mc2-5 subchondral trabecular bone. Results demonstrate that whereas DA values only separate Pongo from African apes (Pan troglodytes, Pan paniscus, Gorilla gorilla), RBV/TV distribution varies with the predicted loading of the metacarpophalangeal (McP) joints during locomotor behaviours in each species. Gorilla exhibits a relatively dorsal distribution of RBV/TV consistent with habitual hyper-extension of the McP joints during knuckle-walking, whereas Pongo has a palmar distribution consistent with flexed McP joints used to grasp arboreal substrates. Both Pan species possess a disto-dorsal distribution of RBV/TV, compatible with multiple hand postures associated with a more varied locomotor regime. Further inter-ray comparisons reveal RBV/TV patterns consistent with varied knuckle-walking postures in Pan species in contrast to higher RBV/TV values toward the midline of the hand in Mc2 and Mc5 of Gorilla, consistent with habitual palm-back knuckle-walking. These patterns of trabecular bone distribution and structure reflect different behavioural signals that could be useful for determining the behaviours of fossil hominins.

Neandertals disappeared from the fossil record around 40,000 bp, after a demographic history of small and isolated groups with high but variable levels of inbreeding, and episodes of interbreeding with other Paleolithic hominins. It is reasonable to expect that high levels of endogamy could be expressed in the skeleton of at least some Neandertal groups. Genetic studies indicate that the 13 individuals from the site of El Sidrón, Spain, dated around 49,000 bp, constituted a closely related kin group, making these Neandertals an appropriate case study for the observation of skeletal signs of inbreeding. We present the complete study of the 1674 identified skeletal specimens from El Sidrón. Altogether, 17 congenital anomalies were observed (narrowing of the internal nasal fossa, retained deciduous canine, clefts of the first cervical vertebra, unilateral hypoplasia of the second cervical vertebra, clefting of the twelfth thoracic vertebra, diminutive thoracic or lumbar rib, os centrale carpi and bipartite scaphoid, tripartite patella, left foot anomaly and cuboid-navicular coalition), with at least four individuals presenting congenital conditions (clefts of the first cervical vertebra). At 49,000 years ago, the Neandertals from El Sidrón, with genetic and skeletal evidence of inbreeding, could be representative of the beginning of the demographic collapse of this hominin phenotype., This research was supported by a grant from the Ministerio de Economía y Competitividad of Spain (CGL2016-75-109-P and Convenio Principado de Asturias–Universidad de Oviedo CN-09–084). L.R. was founded by the Juan de la Cierva program (Spanish Government) and thanks Asier Gómez-Olivencia for providing Neandertal and modern human metric data for the second cervical vertebra and for the discussion of several aspects of the hominin cervical spine. A.E. is founded by the Juan de la Cierva program (Spanish Government). Y.Q. received support from Ministerio de Educación y Ciencia Project N° CGL2015-65387-C3-2-P, MINECO-FEDER and a Predoctoral Grant, University of Burgos. T.L.K is funded by the HP7 European Research Council Starting Grant #336301.

Previously, a micro-finite element (micro-FE)-based inverse remodelling method was presented in the literature that reconstructs the loading history of a bone based on its architecture alone. Despite promising preliminary results, it remains unclear whether this method is sensitive enough to detect differences of bone loading related to pathologies or habitual activities. The goal of this study was to test the sensitivity of the inverse remodelling method by predicting joint loading histories of metacarpal bones of species with similar anatomy but clearly distinct habitual hand use. Three groups of habitual hand use were defined using the most representative primate species: manipulation (human), suspensory locomotion (orangutan), and knuckle-walking locomotion (bonobo, chimpanzee, gorilla). Nine to ten micro-computed tomography scans of each species (\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$n=48$$\end{document}n=48 in total) were used to create micro-FE models of the metacarpal head region. The most probable joint loading history was predicted by optimally scaling six load cases representing joint postures ranging from \documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$-\,75^{\circ }$$\end{document}-75∘ (extension) to \documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$+\,75^{\circ }$$\end{document}+75∘ (flexion). Predicted mean joint load directions were significantly different between knuckle-walking and non-knuckle-walking groups (\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$p

Although much is known about the terrestrial locomotion of great apes, their arboreal locomotion has been studied less extensively. This study investigates arboreal locomotion in bonobos (Pan paniscus), focusing on the gait characteristics and spatio-temporal variables associated with locomotion on a pole. These features are compared across different substrate inclinations (0°, 30°, 45°, 60°, and 90°), and horizontal quadrupedal walking is compared between an arboreal and a terrestrial substrate. Our results show greater variation in footfall patterns with increasing incline, resulting in more lateral gait sequences. During climbing on arboreal inclines, smaller steps and strides but higher stride frequencies and duty factors are found compared to horizontal arboreal walking. This may facilitate better balance control and dynamic stability on the arboreal substrate. We found no gradual change in spatio-temporal variables with increasing incline; instead, the results for all inclines were clustered together. Bonobos take larger strides at lower stride frequencies and lower duty factors on a horizontal arboreal substrate than on a flat terrestrial substrate. We suggest that these changes are the result of the better grip of the grasping feet on an arboreal substrate. Speed modulation of the spatio-temporal variables is similar across substrate inclinations and between substrate types, suggesting a comparable underlying motor control. Finally, we contrast these variables of arboreal inclined climbing with those of terrestrial bipedal locomotion, and briefly discuss the results with respect to the origin of habitual bipedalism. Am. J. Primatol. 78:1165-1177, 2016. © 2016 Wiley Periodicals, Inc.

Objectives Bonobos and chimpanzees are known to differ in various morphological traits, a dichotomy that is sometimes used as an analogy for evolutionary splits during human evolution. The aim of our study was to measure the forearm length of immature and adult bonobos and adult chimpanzees to assess the extent of age–related changes of forearm length in bonobos and sex–dimorphism in bonobos and chimpanzees. Materials and methods As a proxy of somatic growth we measured forearm length of captive bonobos and chimpanzees ranging in age from 1 to 55 years. Measures were taken from subjects inserting their arms into a transparent Plexiglas® tube, a novel technique facilitating repeated measures of nonanesthetized apes in captivity. Results Measures from adult females (>12 years) showed that bonobos exceed chimpanzees in terms of forearm length and that sexual dimorphism in forearm length is pronounced in chimpanzees, but not in bonobos. Forearm length increased significantly with chronological age in bonobos. Validation tests revealed that the device generates useful data on morphometric dimensions. Discussion In most primates, sexual dimorphism in body size is male-biased and the differences in forearm length in chimpanzees follow this trend. Given that males of the two species did not differ in forearm length, the absence/presence of sexual dimorphism of this trait must be due to differences in somatic growth in females. Our novel method offers an alternative to obtain morphometric measures and facilitates longitudinal studies on somatic growth. Am J Phys Anthropol, 2016. © 2016 Wiley Periodicals, Inc.

Background Musculoskeletal and finite element modelling are often used to predict joint loading and bone strength within the human hand, but there is a lack of in vitro evidence of the force and strain experienced by hand bones. Methods This study presents a novel experimental setup that allows the positioning of a cadaveric digit in a variety of postures with the measurement of force and strain experienced by the third metacarpal. The setup allows for the measurement of fingertip force as well. We tested this experimental setup using three cadaveric human third digits in which the flexor tendons were loaded in two tendon pathways: (1) parallel to the metacarpal bone shaft, with bowstringing; (2) a semi-physiological condition in which the tendons were positioned closer to the bone shaft. Results There is substantial variation in metacarpal net force, metacarpal strain and fingertip force between the two tendon pathways. The net force acting on the metacarpal bone is oriented palmarly in the parallel tendon condition, causing tension along the dorsum of the metacarpal shaft, while the force increases and is oriented dorsally in the semi-physiological condition, causing compression of the dorsal metacarpal shaft. Fingertip force is also greater in the semi-physiological condition, implying a more efficient grip function. Inter-individual variation is observed in the radioulnar orientation of the force experienced by the metacarpal bone, the fingertip force, and the strain patterns on the metacarpal shaft. Conclusion This study demonstrates a new method for measuring force and strain experienced by the metacarpal, and fingertip force in cadaveric digits that can, in turn, inform computation models. Inter-individual variation in loads experienced by the third digit suggest that there are differences in joint contact and/or internal bone structure across individuals that are important to consider in clinical and evolutionary contexts.

Objectives Trabecular bone structure is known to be influenced by joint loading during life. However, many additional variables have the potential to contribute to trabecular bone structure of an adult individual, including age, sex, body size, genetics, and overall activity level. There is little research into intraspecific variability in trabecular bone and ontogeny of trabecular bone structure, especially in nonhuman primates. Materials and methods This study investigates trabecular structure in adult and immature chimpanzees from a single population using high-resolution microcomputed tomographic scans of the proximal humerus, proximal femur, and distal tibia. Trabecular bone volume fraction (BV/TV), trabecular thickness (Tb.Th), trabecular number (Tb.N), trabecular spacing (Tb.Sp), and degree of anisotropy (DA) were quantified in specific regions of adult and immature chimpanzees, and color maps were generated to visualize the distribution of BV/TV throughout the joint in the metaphysis of immature specimens. Results The results demonstrate that variability in adult trabecular structure cannot be explained by sex or body size. During ontogeny, there is a general increase in trabecular BV/TV and Tb.Th with age, and ratios of trabecular parameters between the fore- and hindlimb may be consistent with locomotor transitions during ontogeny. Discussion Variation in trabecular morphology among adult individuals is not related to sex or body size, and the factors contributing to intraspecific variability, such as overall activity levels and genetic differences, require further investigation. Trabecular ontogeny in chimpanzees differs from humans in some respects, most notably the absence of a high BV/TV at birth.

Understanding the timing and character of the expansion of Homo sapiens out of Africa is critical for inferring the colonization and admixture processes that underpin global population history. It has been argued that dispersal out of Africa had an early phase, particularly ~130–90 thousand years ago (ka), that reached only the East Mediterranean Levant, and a later phase, ~60–50 ka, that extended across the diverse environments of Eurasia to Sahul. However, recent findings from East Asia and Sahul challenge this model. Here we show that H. sapiens was in the Arabian Peninsula before 85 ka. We describe the Al Wusta-1 (AW-1) intermediate phalanx from the site of Al Wusta in the Nefud desert, Saudi Arabia. AW-1 is the oldest directly dated fossil of our species outside Africa and the Levant. The palaeoenvironmental context of Al Wusta demonstrates that H. sapiens using Middle Palaeolithic stone tools dispersed into Arabia during a phase of increased precipitation driven by orbital forcing, in association with a primarily African fauna. A Bayesian model incorporating independent chronometric age estimates indicates a chronology for Al Wusta of ~95–86 ka, which we correlate with a humid episode in the later part of Marine Isotope Stage 5 known from various regional records. Al Wusta shows that early dispersals were more spatially and temporally extensive than previously thought. Early H. sapiens dispersals out of Africa were not limited to winter rainfall-fed Levantine Mediterranean woodlands immediately adjacent to Africa, but extended deep into the semi-arid grasslands of Arabia, facilitated by periods of enhanced monsoonal rainfall.

Aspects of trabecular bone architecture are thought to reflect regional loading of the skeleton, and thus differ between primate taxa with different locomotor and postural modes. However, there are several systemic factors that affect bone structure that could contribute to, or be the primary factor determining, interspecific differences in bone structure. These systemic factors include differences in genetic regulation, sensitivity to loading, hormone levels, diet, and activity levels. Improved understanding of inter-/intraspecific variability, and variability across the skeleton of an individual, is required to interpret properly potential functional signals present within trabecular structure. Using a whole-region method of analysis, we investigated trabecular structure throughout the skeleton of humans and chimpanzees. Trabecular bone volume fraction (BV/TV), degree of anisotropy (DA) and trabecular thickness (Tb.Th) were quantified from high resolution micro-computed tomographic scans of the humeral and femoral head, third metacarpal and third metatarsal head, distal tibia, talus and first thoracic vertebra. We found that BV/TV is, in most anatomical sites, significantly higher in chimpanzees than in humans, suggesting a systemic difference in trabecular structure unrelated to local loading regime. Differences in BV/TV between the forelimb and hindlimb did not clearly reflect differences in locomotor loading in the study taxa. There were no clear systemic differences between the taxa in DA and, as such, this parameter might reflect function and relate to differences in joint loading. This systemic approach reveals both the pattern of variability across the skeleton and between taxa, and helps identify those features of trabecular structure that may relate to joint function.

Evolution of the human hand has undergone a transition from use during locomotion to use primarily for manipulation. Previous comparative morphological and biomechanical studies have focused on potential changes in manipulative abilities during human hand evolution, but few have focused on functional signals for arboreal locomotion. Here, we provide this comparative context though the first analysis of hand loading in captive bonobos during arboreal locomotion. We quantify pressure experienced by the fingers, palm and thumb in bonobos during vertical locomotion, suspension and arboreal knuckle-walking. Results show that pressure experienced by the fingers is significantly higher during knuckle-walking compared with similar pressures experienced by the fingers and palm during suspensory and vertical locomotion. Peak pressure is most often experienced at or around the third digit in all locomotor modes. Pressure quantified for the thumb is either very low or absent, despite the thumb making contact with the substrate during all suspensory and vertical locomotor trials. Unlike chimpanzees, the bonobos do not show a rolling pattern of digit contact with the substrate during arboreal knuckle-walking but, instead, digits 3 and 4 typically touch down first and digit 5 almost always made contact with the substrate. These results have implications for interpreting extant and fossilised hand morphology; we expect bonobo (and chimpanzee) bony morphology to primarily reflect the biomechanical loading of knuckle-walking, while functional signals for arboreal locomotion in fossil hominins are most likely to appear in the fingers, particularly digit 3, and least likely to appear in the morphology of the thumb.

Getting a grip The evolution of the hand—particularly the opposable thumb—was key to the success of early humans. Without a precise grip, involving forceful opposition of thumb with fingers, tool technology could not have emerged. Skinner et al. analyzed the internal bone structure of Pliocene Australopithecus hands, dated at 3.2 million years old. Internal bone structure reveals the patterns and directions of forces operating on the hand, providing clues to the kinds of activities performed. Modern human-like hand postures consistent with the habitual use of tools appeared about half a million years earlier than the first archaeological evidence of stone tools. Science , this issue p. 395

Hand bone morphology is regularly used to link particular hominin species with behaviors relevant to cognitive/technological progress. Debates about the functional significance of differing hominin hand bone morphologies tend to rely on establishing phylogenetic relationships and/or inferring behavior from epigenetic variation arising from mechanical loading and adaptive bone modeling. Most research focuses on variation in cortical bone structure, but additional information about hand function may be provided through the analysis of internal trabecular structure. While primate hand bone trabecular structure is known to vary in ways that are consistent with expected joint loading differences during manipulation and locomotion, no study exists that has documented this variation across the numerous bones of the hand. We quantify the trabecular structure in 22 bones of the human hand (early/extant modern Homo sapiens) and compare structural variation between two groups associated with post-agricultural/industrial (post-Neolithic) and foraging/hunter-gatherer (forager) subsistence strategies. We (1) establish trabecular bone volume fraction (BV/TV), modulus (E), degree of anisotropy (DA), mean trabecular thickness (Tb.Th) and spacing (Tb.Sp); (2) visualize the average distribution of site-specific BV/TV for each bone; and (3) examine if the variation in trabecular structure is consistent with expected joint loading differences among the regions of the hand and between the groups. Results indicate similar distributions of trabecular bone in both groups, with those of the forager sample presenting higher BV/TV, E, and lower DA, suggesting greater and more variable loading during manipulation. We find indications of higher loading along the ulnar side of the forager sample hand, with high site-specific BV/TV distributions among the carpals that are suggestive of high loading while the wrist moves through the ‘dart-thrower's’ motion. These results support the use of trabecular structure to infer behavior and have direct implications for refining our understanding of human hand evolution and fossil hominin hand use.

Evolution of the human hand has undergone a transition from use during locomotion to use primarily for manipulation. Previous comparative morphological and biomechanical studies have focused on potential changes in manipulative abilities during human hand evolution, but few have focused on functional signals for arboreal locomotion. Here, we provide this comparative context though the first analysis of hand loading in captive bonobos during arboreal locomotion. We quantify pressure experienced by the fingers, palm and thumb in bonobos during vertical locomotion, suspension and arboreal knuckle-walking. The results show that pressure experienced by the fingers is significantly higher during knuckle-walking compared with similar pressures experienced by the fingers and palm during suspensory and vertical locomotion. Peak pressure is most often experienced at or around the third digit in all locomotor modes. Pressure quantified for the thumb is either very low or absent, despite the thumb making contact with the substrate during all suspensory and vertical locomotor trials. Unlike chimpanzees, bonobos do not show a rolling pattern of digit contact with the substrate during arboreal knuckle-walking - instead, we found that digits 3 and 4 typically touch down first and digit 5 almost always made contact with the substrate. These results have implications for interpreting extant and fossilized hand morphology; we expect bonobo (and chimpanzee) bony morphology to primarily reflect the biomechanical loading of knuckle-walking, while functional signals for arboreal locomotion in fossil hominins are most likely to appear in the fingers, particularly digit 3, and least likely to appear in the morphology of the thumb.

It is widely agreed that biomechanical stresses imposed by stone tool behaviors influenced the evolution of the human hand. Though archaeological evidence suggests that early hominins participated in a variety of tool behaviors, it is unlikely that all behaviors equally influenced modern human hand anatomy. It is more probable that a behavior's likelihood of exerting a selective pressure was a weighted function of the magnitude of stresses associated with that behavior, the benefits received from it, and the amount of time spent performing it. Based on this premise, we focused on the first part of that equation and evaluated magnitudes of stresses associated with stone tool behaviors thought to have been commonly practiced by early hominins, to determine which placed the greatest loads on the digits. Manual pressure data were gathered from 39 human subjects using a Novel Pliance® manual pressure system while they participated in multiple Plio-Pleistocene tool behaviors: nut-cracking, marrow acquisition with a hammerstone, flake production with a hammerstone, and handaxe and flake use. Manual pressure distributions varied significantly according to behavior, though there was a tendency for regions of the hand subject to the lowest pressures (e.g., proximal phalanges) to be affected less by behavior type. Hammerstone use during marrow acquisition and flake production consistently placed the greatest loads on the digits collectively, on each digit and on each phalanx. Our results suggest that, based solely on the magnitudes of stresses, hammerstone use during marrow acquisition and flake production are the most likely of the assessed behaviors to have influenced the anatomical and functional evolution of the human hand.

The Rising Star cave system has produced abundant fossil hominin remains within the Dinaledi Chamber, representing a minimum of 15 individuals attributed to Homo naledi. Further exploration led to the discovery of hominin material, now comprising 131 hominin specimens, within a second chamber, the Lesedi Chamber. The Lesedi Chamber is far separated from the Dinaledi Chamber within the Rising Star cave system, and represents a second depositional context for hominin remains. In each of three collection areas within the Lesedi Chamber, diagnostic skeletal material allows a clear attribution to H. naledi. Both adult and immature material is present. The hominin remains represent at least three individuals based upon duplication of elements, but more individuals are likely present based upon the spatial context. The most significant specimen is the near-complete cranium of a large individual, designated LES1, with an endocranial volume of approximately 610 ml and associated postcranial remains. The Lesedi Chamber skeletal sample extends our knowledge of the morphology and variation of H. naledi, and evidence of H. naledi from both recovery localities shows a consistent pattern of differentiation from other hominin species. DOI: http://dx.doi.org/10.7554/eLife.24232.001, eLife digest Species of ancient humans and the extinct relatives of our ancestors are typically described from a limited number of fossils. However, this was not the case with Homo naledi. More than 1500 fossils representing at least 15 individuals of this species were unearthed from the Rising Star cave system in South Africa between 2013 and 2014. Found deep underground in the Dinaledi Chamber, the H. naledi fossils are the largest collection of a single species of an ancient human-relative discovered in Africa. After the discovery was reported, a number of questions still remained. These questions included: why were so many fossils from a single species found at the one site, and how did they come to rest so far into the cave system? Possible explanations such as H. naledi living in the cave or being washed in by a flood were considered but ruled out. Instead, the evidence was largely consistent with intact bodies being deliberately disposed of in the cave and then decomposing. Now, Hawks et al. – who include many of the researchers who were involved in the discovery of H. naledi – report that yet more H. naledi fossils have been unearthed from a second chamber in the Rising Star cave system, the Lesedi Chamber. The chamber is 30 meters below the surface and there is no direct route between it and the Dinaledi Chamber. Again, the evidence is most consistent with the bodies arriving intact into the chamber, and there were no signs that the remains had been exposed to the surface environment. Also like the Dinaledi Chamber, no remains of other ancient humans or their relatives were found in the Lesedi Chamber. In total, 133 fossils of H. naledi have been found in this second chamber representing at least three individuals: two adults and a juvenile. However, and as Hawks et al. point out, only a small volume of the chamber has been excavated so far, and so there are likely more fossils still to be found. The fossils in the Lesedi Chamber are similar to those found before but include intact examples of bones, like the collarbone, that were previously known only from fragments. Perhaps the most impressive among the new fossils is a relatively complete skull that is part of a partial skeleton. The skull could have housed a brain that was 9% larger than the maximum estimate calculated from the previous H. naledi fossils. Though these new fossils provide us with yet more information about H. naledi, some questions still remain unanswered – the material from the Lesedi Chamber is undated, for example. However, a related study by Dirks et al. does give an estimate for the age of the fossils from the Dinaledi Chamber, while Berger et al. provide an explanation for why this date might be much younger than was previously predicted. DOI: http://dx.doi.org/10.7554/eLife.24232.002