Southern African hominin fossils traditionally attributed to Paranthropus robustus and Australopithecus africanus are differentiated from each other by their dentition and cranial architecture, but their postcranial anatomy has typically been regarded functionally as broadly similar (i.e., terrestrial bipedalism with some degree of arboreal locomotion). Testing the hypothesis of a similar locomotor repertoire between these two taxa has been complicated by a lack of postcranial fossils attributable to P. robustus. Here, we detail our comparative examination of the internal bone anatomy of a recently described c. 1.8 Ma P. robustus articulating femur and tibia, which suggests distinct patterns of joint loading and locomotor behavior. Our analysis of cortical bone in the hip and trabecular bone in the ankle, knee, and hip joints suggests that P. robustus habitually adopted a high frequency of ankle, knee, and hip flexion, as required for climbing, while Australopithecus from a c. 3.4 Ma level at the site of Sterkfontein displays a more modern human-like structural pattern across the lower limb joints consistent with comparatively more frequent terrestrial bipedalism. These results reveal that geologically younger P. robustus likely used arboreality more frequently than older Sterkfontein Australopithecus. Together with differences in masticatory behavior, this line of evidence indicates that these two hominin taxa occupied distinct ecological niches.

OBJECTIVES:Extant humans and non-human apes are characterized by diverse locomotor and postural behaviors, resulting in different joint loading patterns. These behaviors influence trabecular bone, which responds to mechanical loading and joint posture. While prior studies have examined femoral trabecular structure, this study is the first to assess trabecular architecture in the proximal tibia across extant hominoids to evaluate how differences in joint use and posture are reflected in tibial morphology. MATERIALS AND METHODS:We analyze trabecular structure in the proximal tibiae of Homo sapiens (n = 25), Gorilla (n = 13), Pan troglodytes (n = 15) and Pongo sp. (n = 7). Each tibia was scanned using high-resolution microtomography, and cortical and trabecular bone were isolated from each other. Canonical holistic morphometric analysis was used to quantify trabecular bone volume fraction and degree of anisotropy. Spatial distributions of these variables were compared across taxa using principal component analysis, and group differences were assessed with multivariate analysis of variance and pairwise tests. RESULTS:Results show that trabecular bone volume and anisotropy reflect habitual knee posture: extended in bipedal Homo, and flexed in non-human apes. In Gorilla, males exhibit more extended knee use than females, while no significant sex differences were observed in Homo or Pan (sex differences in Pongo were not tested due to sample limitations). DISCUSSION:We demonstrate that the trabecular structure of the proximal tibia is consistent with habitual locomotor loading in extant hominids, which provides the comparative context to interpret knee posture, biomechanical loading, and predominant locomotor behaviors in fossil hominids.

The evolution of the human hand is marked by a transition from a hand primarily used for locomotion to one primarily used for dexterous manipulation. The hand skeletons of Plio-Pleistocene hominins have different mosaics of human-like features associated with enhanced dexterity and ape-like features associated with locomotor hand use. However, the functional relevance of the ape-like features is debated, particularly due to a lack of complete and associated hand remains. Here, we investigate the internal phalangeal cortical structure of the nearly complete Australopithecus sediba MH2 hand and Homo naledi hand 1 to provide both insight into the manual behaviors of these fossil hominins and functional clarity regarding the mosaic features found within their hands. The phalangeal cortical structure demonstrates diversity in Plio-Pleistocene hand use, with A. sediba and H. naledi each indicating different dexterous abilities and different climbing strategies, supporting the functional importance of the ape-like features.

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. This study provided a holistic approach of trabecular bone architecture within the hominid distal femur. Trabecular structure in Homo reflects habitual use of extended knee postures during bipedalism, habitual use of flexed knee posture during terrestrial and arboreal locomotion in African apes, and the highly mobile knee joint in Pongo. This study offers a comparative sample of trabecular structure in the hominoid distal femur and can contribute to future studies of locomotion in extinct taxa.image

The “gold standard” for the assessment of trabecular bone structure is high-resolution micro-CT. In this technical note, we test the influence of initial scan resolution and post hoc downsampling on the quantitative and qualitative analysis of trabecular bone in a Gorilla tibia. We analyzed trabecular morphology in the right distal tibia of one Gorilla gorilla individual to investigate the impact of variation in voxel size on measured trabecular variables. For each version of the micro-CT volume, trabecular bone was segmented using the medical image analysis method. Holistic morphometric analysis was then used to analyze bone volume (BV/TV), anisotropy (DA), trabecular thickness (Tb.Th), spacing (Tb.Sp), and number (Tb.N). Increasing voxel size during initial scanning was found to have a strong impact on DA and Tb.Th measures, while BV/TV, Tb.Sp, and Tb.N were found to be less sensitive to variations in initial scan resolution. All tested parameters were not substantially influenced by downsampling up to 90 μm resolution. Color maps of BV/TV and DA also retained their distribution up to 90 μm. This study is the first to examine the effect of variation in micro-CT voxel size on the analysis of trabecular bone structure using whole epiphysis approaches. Our results indicate that microstructural variables may be measured for most trabecular parameters up to a voxel size of 90 μm for both scan and downsampled resolutions. Moreover, if only BV/TV, Tb.Sp or Tb.N is measured, even larger voxel sizes might be used without substantially affecting the results.

OBJECTIVES:This research examines whether the distribution of trabecular bone in the proximal capitates of extant hominids, as well as several fossil hominin taxa, is associated with the oblique path of the midcarpal joint known as the dart-thrower's motion (DTM). MATERIALS AND METHODS:We analyzed proximal capitates from extant (Pongo n = 12; Gorilla n = 11; Pan n = 10; fossil and recent Homo sapiens n = 29) and extinct (Australopithecus sediba n = 2; Homo naledi n = 1; Homo floresiensis n = 2; Neandertals n = 3) hominids using a new canonical holistic morphometric analysis, which quantifies and visualizes the distribution of trabecular bone using relative bone volume as a fraction of total volume (rBV/TV). RESULTS:Homo sapiens and Neandertals had a continuous band of high rBV/TV that extended across the scaphoid, lunate, and hamate subarticular regions, but other fossil hominins and extant great apes did not. A. sediba expressed a distinct combination of human-like and Pan-like rBV/TV distribution. Both H. floresiensis and H. naledi had high rBV/TV on the ulnar-side of the capitate but low rBV/TV on the radial-side. CONCLUSION:The proximal capitates of H. sapiens and Neandertals share a distinctive distribution of trabecular bone that suggests that these two species of Homo regularly load(ed) their midcarpal joints along the full extent of the oblique path of the DTM. The observed pattern in A. sediba suggests that human-like stress at the capito-scaphoid articular surface was combined with Pan-like wrist postures, whereas the patterns in H. floresiensis and H. naledi suggest their midcarpal joints were loaded differently from that of H. sapiens and Neandertals.

OBJECTIVES Recent studies have associated subarticular trabecular bone distribution in the extant hominid first metacarpal (Mc1) with observed thumb use, to infer fossil hominin thumb use. Here, we analyze the entire Mc1 to test for interspecific differences in: (1) the absolute volume of trabecular volume fraction, (2) the distribution of the deeper trabecular network, and (3) the distribution of trabeculae in the medullary cavity, especially beneath the Mc1 disto-radial flange. MATERIALS AND METHODS Trabecular bone was imaged using micro-computed tomography in a sample of Homo sapiens (n = 11), Pan paniscus (n = 10), Pan troglodytes (n = 11), Gorilla gorilla (n = 10) and Pongo sp., (n = 7). Using Canonical Holistic Morphometric Analysis (cHMA), we tested for interspecific differences in the trabecular bone volume fraction (BV/TV) and its relative distribution (rBV/TV) throughout the Mc1, including within the head, medullary cavity, and base. RESULTS P. paniscus had the highest, and H. sapiens the lowest, BV/TV relative to other species. rBV/TV distribution statistically distinguished the radial concentrations and lack of medullary trabecular bone in the H. sapiens Mc1 from all other hominids. H. sapiens and, to a lesser extent, G. gorilla also had a significantly higher trabecular volume beneath the disto-radial flange relative to other hominids. DISCUSSION These results are consistent with differences in observed thumb use in these species and may also reflect systemic differences in bone volume fraction. The trabecular bone extension into the medullary cavity and concentrations beneath the disto-radial flange may represent crucial biomechanical signals that will aid in the inference of fossil hominin thumb use.

OBJECTIVES:Analyses of external bone shape using geometric morphometrics (GM) and cross-sectional geometry (CSG) are frequently employed to investigate bone structural variation and reconstruct activity in the past. However, the association between these methods has not been thoroughly investigated. Here, we analyze whole bone shape and CSG variation of metacarpals 1-5 and test covariation between them. MATERIALS AND METHODS:We analyzed external metacarpal shape using GM and CSG of the diaphysis at three locations in metacarpals 1-5. The study sample includes three modern human groups: crew from the shipwrecked Mary Rose (n = 35 metacarpals), a Pre-industrial group (n = 50), and a Post-industrial group (n = 31). We tested group differences in metacarpal shape and CSG, as well as correlations between these two aspects of metacarpal bone structure. RESULTS:GM analysis demonstrated metacarpus external shape variation is predominately related to changes in diaphyseal width and articular surface size. Differences in external shape were found between the non-pollical metacarpals of the Mary Rose and Pre-industrial groups and between the third metacarpals of the Pre- and Post-industrial groups. CSG results suggest the Mary Rose and Post-industrial groups have stronger metacarpals than the Pre-industrial group. Correlating CSG and external shape showed significant relationships between increasing external robusticity and biomechanical strength across non-pollical metacarpals (r: 0.815-0.535; p ≤ 0.05). DISCUSSION:Differences in metacarpal cortical structure and external shape between human groups suggest differences in the type and frequency of manual activities. Combining these results with studies of entheses and kinematics of the hand will improve reconstructions of manual behavior in the past.

OBJECTIVES The shape of the trapezium and first metacarpal (Mc1) markedly influence thumb mobility, strength, and the manual abilities of extant hominids. Previous research has typically focused solely on trapezium-Mc1 joint shape. Here we investigate how morphological integration and shape covariation between the entire trapezium (articular and non-articular surfaces) and the entire Mc1 reflect known differences in thumb use in extant hominids. MATERIALS AND METHODS We analyzed shape covariation in associated trapezia and Mc1s across a large, diverse sample of Homo sapiens (n = 40 individuals) and other extant hominids (Pan troglodytes, n = 16; Pan paniscus, n = 13; Gorilla gorilla gorilla, n = 27; Gorilla beringei, n = 6; Pongo pygmaeus, n = 14; Pongo abelii, n = 9) using a 3D geometric morphometric approach. We tested for interspecific significant differences in degree of morphological integration and patterns of shape covariation between the entire trapezium and Mc1, as well as within the trapezium-Mc1 joint specifically. RESULTS Significant morphological integration was only found in the trapezium-Mc1 joint of H. sapiens and G. g. gorilla. Each genus showed a specific pattern of shape covariation between the entire trapezium and Mc1 that was consistent with different intercarpal and carpometacarpal joint postures. DISCUSSION Our results are consistent with known differences in habitual thumb use, including a more abducted thumb during forceful precision grips in H. sapiens and a more adducted thumb in other hominids used for diverse grips. These results will help to infer thumb use in fossil hominins.

Bone is a remarkable, living tissue that functionally adapts to external loading. Therefore, bone shape and internal structure carry information relevant to many disciplines, including medicine, forensic science, and anthropology. However, morphometric comparisons of homologous regions across different individuals or groups are still challenging. In this study, two methods were combined to quantify such differences: (1) Holistic morphometric analysis (HMA) was used to quantify morphometric values in each bone, (2) which could then be mapped to a volumetric mesh of a canonical bone created by a statistical free-form deformation model (SDM). Required parameters for this canonical holistic morphometric analysis (cHMA) method were identified and the robustness of the method was evaluated. The robustness studies showed that the SDM converged after one to two iterations, had only a marginal bias towards the chosen starting image, and could handle large shape differences seen in bones of different species. Case studies were performed on metacarpal bones and proximal femora of different primate species to confirm prior study results. The differences between species could be visualised and statistically analysed in both case studies. cHMA provides a framework for performing quantitative comparisons of different morphometric quantities across individuals or groups. These comparisons facilitate investigation of the relationship between spatial morphometric variations and function or pathology, or both.

The human lineage is marked by a transition in hand use, from locomotion towards increasingly dexterous manipulation, concomitant with bipedalism. The forceful precision grips used by modern humans probably evolved in the context of tool manufacture and use, but when and how many times hominin hands became principally manipulative remains unresolved. We analyse metacarpal trabecular and cortical bone, which provide insight into behaviour during an individual’s life, to demonstrate previously unrecognized diversity in hominin hand use. The metacarpals of the palm in Australopithecus sediba have trabecular morphology most like orangutans and consistent with locomotor power-grasping with the fingers, while that of the thumb is consistent with human-like manipulation. This internal morphology is the first record of behaviour consistent with a hominin that used its hand for both arboreal locomotion and human-like manipulation. This hand use is distinct from other fossil hominins in this study, including A. afarensis and A. africanus .

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

Much research has debated the technological abilities of Neanderthals relative to those of early modern humans, with a particular focus on subtle differences in thumb morphology and how this may reflect differences in manipulative behaviors in these two species. Here, we provide a novel perspective on this debate through a 3D geometric morphometric analysis of shape covariation between the trapezial and proximal first metacarpal articular surfaces of Neanderthals ( Homo neanderthalensis ) in comparison to early and recent humans ( Homo sapiens ). Results show a distinct pattern of shape covariation in Neanderthals, consistent with more extended and adducted thumb postures that may reflect habitual use of grips commonly used for hafted tools. Both Neanderthals and recent humans demonstrate high intraspecific variation in shape covariation. This intraspecific variation is likely the result of genetic and/or developmental differences, but may also reflect, in part, differing functional requirements imposed by the use of varied tool-kits. These results underscore the importance of holistic joint shape analysis for understanding the functional capabilities and evolution of the modern human thumb.

Bipedalism is a defining trait of the hominin lineage, associated with a transition from a more arboreal to a more terrestrial environment. While there is debate about when modern human-like bipedalism first appeared in hominins, all known South African hominins show morphological adaptations to bipedalism, suggesting that this was their predominant mode of locomotion. Here we present evidence that hominins preserved in the Sterkfontein Caves practiced two different locomotor repertoires. The trabecular structure of a proximal femur (StW 522) attributed toAustralopithecus africanusexhibits a modern human-like bipedal locomotor pattern, while that of a geologically younger specimen (StW 311) attributed to eitherHomosp. orParanthropus robustusexhibits a pattern more similar to nonhuman apes, potentially suggesting regular bouts of both climbing and terrestrial bipedalism. Our results demonstrate distinct morphological differences, linked to behavioral differences betweenAustralopithecusand later hominins in South Africa and contribute to the increasing evidence of locomotor diversity within the hominin clade.