What do brain endocasts tell us? A comparative analysis of the accuracy of sulcal identification by experts and perspectives in palaeoanthropology.

Palaeoneurology is a complex field as the object of study, the brain, does not fossilize. Studies rely therefore on the (brain) endocranial cast (often named endocast), the only available and reliable proxy for brain shape, size and details of surface. However, researchers debate whether or not specific marks found on endocasts correspond reliably to particular sulci and/or gyri of the brain that were imprinted in the braincase. The aim of this study is to measure the accuracy of sulcal identification through an experiment that reproduces the conditions that palaeoneurologists face when working with hominin endocasts. We asked 14 experts to manually identify well-known foldings in a proxy endocast that was obtained from an MRI of an actual in vivo Homo sapiens head. We observe clear differences in the results when comparing the non-corrected labels (the original labels proposed by each expert) with the corrected labels. This result illustrates that trying to reconstruct a sulcus following the very general known shape/position in the literature or from a mean specimen may induce a bias when looking at an endocast and trying to follow the marks observed there. We also observe that the identification of sulci appears to be better in the lower part of the endocast compared to the upper part. The results concerning specific anatomical traits have implications for highly debated topics in palaeoanthropology. Endocranial description of fossil specimens should in the future consider the variation in position and shape of sulci in addition to using models of mean brain shape. Moreover, it is clear from this study that researchers can perceive sulcal imprints with reasonably high accuracy, but their correct identification and labelling remains a challenge, particularly when dealing with extinct species for which we lack direct knowledge of the brain.

Mechanical compensation in the evolution of the early hominin feeding apparatus.

Australopiths, a group of hominins from the Plio-Pleistocene of Africa, are characterized by derived traits in their crania hypothesized to strengthen the facial skeleton against feeding loads and increase the efficiency of bite force production. The crania of robust australopiths are further thought to be stronger and more efficient than those of gracile australopiths. Results of prior mechanical analyses have been broadly consistent with this hypothesis, but here we show that the predictions of the hypothesis with respect to mechanical strength are not met: some gracile australopith crania are as strong as that of a robust australopith, and the strength of gracile australopith crania overlaps substantially with that of chimpanzee crania. We hypothesize that the evolution of cranial traits that increased the efficiency of bite force production in australopiths may have simultaneously weakened the face, leading to the compensatory evolution of additional traits that reinforced the facial skeleton. The evolution of facial form in early hominins can therefore be thought of as an interplay between the need to increase the efficiency of bite force production and the need to maintain the structural integrity of the face.

Trabecular bone properties in the ilium of the Middle Paleolithic/Middle Stone Age Border Cave 3 Homo sapiens infant and the onset of independent gait.

The Border Cave 3 (BC3) infant skeleton has been understudied, despite its importance as an example of a well-preserved and fairly complete immature skeleton of early Homo sapiens which potentially provides a rare window into various aspects of ontogenetic development, including locomotor activity (e.g., timing of gait events). Trabecular structure in the BC3 ilium was evaluated to investigate whether it matches that of an equivalently aged infant from a postindustrialized society. Microcomputed tomography (µCT) scans were acquired from the BC3 infant and from an ontogenetic series of 25 postindustrial infants that were divided into three age classes (ACs) ranging from neonates to toddlers (<36 months). All ilia were qualitatively compared and then digitally subdivided into 10 volumes of interest (VOIs) based on anatomical reference points. The VOIs were quantified and ontogenetic differences in trabecular structure were statistically evaluated. Across the comparative ontogenetic series, trabecular architectural properties overlapped in all regions. However, trabecular thickness increased significantly after the first year of life. The BC3 infant demonstrated generally similar trabecular structure to that observed in the age-equivalent postindustrial infants (AC2), including relatively strong development of the trabecular chiasma qualitatively. However, some interesting distinctions were observed in BC3, such as low strut thickness compared with infants from the postindustrial sample, that bear further exploration in future studies. Evaluation of only one individual from the Middle Stone Age (MSA), coupled with the relatively small comparative sample, limit our ability to distinguish more meaningful biological differences in trabecular structure throughout ontogeny from idiosyncratic characteristics. Nonetheless, results of this study extend ongoing research on infant locomotor and morphological development to archeological populations in the Middle Stone Age. Further cross-cultural studies consisting of larger comparative postindustrial samples may provide additional information on trabecular structure in the infant ilium during this important developmental timeframe.

Morphological description and evolutionary significance of 300 ka hominin facial bones from Hualongdong, China.

Late Middle Pleistocene hominins in Africa displaying key modern morphologies by 315 ka are claimed as the earliest Homo sapiens. Evolutionary relationships among East Asian hominins appear complex due to a growing fossil record of late Middle Pleistocene hominins from the region, reflecting mosaic morphologies that contribute to a lack of consensus on when and how the transition to modern humans transpired. Newly discovered 300 ka hominin fossils from Hualongdong, China, provide further evidence to clarify these relationships in the region. In this study, facial morphology of the juvenile partial cranium (HLD 6) is described and qualitatively and quantitatively compared with that of other key Early, Middle, and Late Pleistocene hominins from East Asia, Africa, West Asia, and Europe and with a sample of modern humans. Qualitatively, facial morphology of HLD 6 resembles that of Early and Middle Pleistocene hominins from Zhoukoudian, Nanjing, Dali, and Jinniushan in China, as well as others from Java, Africa, and Europe in some of these features (e.g., supraorbital and malar regions), and Late Pleistocene hominins and modern humans from East Asia, Africa, and Europe in other features (e.g., weak prognathism, flat face and features in nasal and hard plate regions). Comparisons of HLD 6 measurements to group means and multivariate analyses support close affinities of HLD 6 to Late Pleistocene hominins and modern humans. Expression of a mosaic morphological pattern in the HLD 6 facial skeleton further complicates evolutionary interpretations of regional morphological diversity in East Asia. The prevalence of modern features in HLD 6 suggests that the hominin population to which HLD 6 belonged may represent the earliest pre-modern humans in East Asia. Thus, the transition from archaic to modern morphology in East Asian hominins may have occurred at least by 300 ka, which is 80,000 to 100,000 years earlier than previously recognized.

The pectoral girdle of StW 573 ('Little Foot') and its implications for shoulder evolution in the Hominina.

The ca. 3.67 Ma adult skeleton known as 'Little Foot' (StW 573), recovered from Sterkfontein Member 2 breccia in the Silberberg Grotto, is remarkable for its morphology and completeness. Preservation of clavicles and scapulae, including essentially complete right-side elements, offers opportunities to assess morphological and functional aspects of a nearly complete Australopithecus pectoral girdle. Here we describe the StW 573 pectoral girdle and offer quantitative comparisons to those of extant hominoids and selected homininans. The StW 573 pectoral girdle combines features intermediate between those of humans and other apes: a long and curved clavicle, suggesting a relatively dorsally positioned scapula; an enlarged and uniquely proportioned supraspinous fossa; a relatively cranially oriented glenoid fossa; and ape-like reinforcement of the axillary margin by a stout ventral bar. StW 573 scapulae are as follows: smaller than those of some homininans (i.e., KSD-VP-1/1 and KNM-ER 47000A), larger than others (i.e., A.L. 288-1, Sts 7, and MH2), and most similar in size to another australopith from Sterkfontein, StW 431. Moreover, StW 573 and StW 431 exhibit similar structural features along their axillary margins and inferior angles. As the StW 573 pectoral girdle (e.g., scapular configuration) has a greater affinity to that of apes-Gorilla in particular-rather than modern humans, we suggest that the StW 573 morphological pattern appears to reflect adaptations to arboreal behaviors, especially those with the hand positioned above the head, more than human-like manipulatory capabilities. When compared with less complete pectoral girdles from middle/late Miocene apes and that of the penecontemporaneous KSD-VP-1/1 (Australopithecus afarensis), and mindful of consensus views on the adaptiveness of arboreal positional behaviors soliciting abducted glenohumeral joints in early Pliocene taxa, we propose that the StW 573 pectoral girdle is a reasonable model for hypothesizing pectoral girdle configuration of the crown hominin last common ancestor.

Unique foot posture in Neanderthals reflects their body mass and high mechanical stress.

Neanderthal foot bone proportions and morphology are mostly indistinguishable from those of Homo sapiens, with the exception of several distinct Neanderthal features in the talus. The biomechanical implications of these distinct talar features remain contentious, fueling debate around the adaptive meaning of this distinctiveness. With the aim of clarifying this controversy, we test phylogenetic and behavioral factors as possible contributors, comparing tali of 10 Neanderthals and 81 H. sapiens (Upper Paleolithic and Holocene hunter-gatherers, agriculturalists, and postindustrial group) along with the Clark Howell talus (Omo, Ethiopia). Variation in external talar structures was assessed through geometric morphometric methods, while bone volume fraction and degree of anisotropy were quantified in a subsample (n = 45). Finally, covariation between point clouds of site-specific trabecular variables and surface landmark coordinates was assessed. Our results show that although Neanderthal talar external and internal morphologies were distinct from those of H. sapiens groups, shape did not significantly covary with either bone volume fraction or degree of anisotropy, suggesting limited covariation between external and internal talar structures. Neanderthal external talar morphology reflects ancestral retentions, along with various adaptations to high levels of mobility correlated to their presumably unshod hunter-gatherer lifestyle. This pairs with their high site-specific trabecular bone volume fraction and anisotropy, suggesting intense and consistently oriented locomotor loading, respectively. Relative to H.sapiens, Neanderthals exhibit differences in the talocrural joint that are potentially attributable to cultural and locomotor behavior dissimilarity, a talonavicular joint that mixes ancestral and functional traits, and a derived subtalar joint that suggests a predisposition for a pronated foot during stance phase. Overall, Neanderthal talar variation is attributable to mobility strategy and phylogenesis, while H. sapiens talar variation results from the same factors plus footwear. Our results suggest that greater Neanderthal body mass and/or higher mechanical stress uniquely led to their habitually pronated foot posture.

Combinations of trabecular and cortical bone properties distinguish various loading modalities between athletes and controls.

OBJECTIVES: Variation in trabecular and cortical bone properties is often used to infer habitual behavior in the past. However, the structures of both types of bone are rarely considered together and may even contradict each other in functional interpretations. We examine trabecular and cortical bone properties in various athletes and sedentary controls to clarify the associations between combinations of cortical and trabecular bone properties and various loading modalities. MATERIALS AND METHODS: We compare trabecular and cortical bone properties using peripheral quantitative computed tomography scans of the tibia between groups of 83 male athletes (running, hockey, swimming, cricket) and sedentary controls using Bayesian multilevel models. We quantify midshaft cortical bone rigidity and area (J, CA), midshaft shape index (Imax/Imin), and mean trabecular bone mineral density (BMD) in the distal tibia. RESULTS: All groups show unique combinations of biomechanical properties. Cortical bone rigidity is high in sports that involve impact loading (cricket, running, hockey) and low in nonimpact loaded swimmers and controls. Runners have more anteroposteriorly elliptical midshafts compared to other groups. Interestingly, all athletes have greater trabecular BMD compared to controls, but do not differ credibly among each other. DISCUSSION: Results suggest that cortical midshaft hypertrophy is associated with impact loading while trabecular BMD is positively associated with both impact and nonimpact loading. Midshaft shape is associated with directionality of loading. Individuals from the different categories overlap substantially, but group means differ credibly, suggesting that nuanced group-level inferences of habitual behavior are possible when combinations of trabecular and cortical bone are analyzed.

Automated resolution independent method for comparing in vivo and dry trabecular bone.

OBJECTIVES: Variation in human trabecular bone morphology can be linked to habitual behavior, but it is difficult to investigate in vivo due to the radiation required at high resolution. Consequently, functional interpretations of trabecular morphology remain inferential. Here we introduce a method to link low- and high-resolution CT data from dry and fresh bone, enabling bone functional adaptation to be studied in vivo and results compared to the fossil and archaeological record. MATERIALS AND METHODS: We examine 51 human dry bone distal tibiae from Nile Valley and UK and two pig tibiae containing soft tissues. We compare low-resolution peripheral quantitative computed tomography (pQCT) parameters and high-resolution micro CT (µCT) in homologous single slices at 4% bone length and compare results to our novel Bone Ratio Predictor (BRP) method. RESULTS: Regression slopes between linear attenuation coefficients of low-resolution pQCT images and bone area/total area (BA/TA) of high-resolution µCT scans differ substantially between geographical subsamples, presumably due to diagenesis. BRP accurately predicts BA/TA (R2 = .97) and eliminates the geographic clustering. BRP accurately estimates BA/TA in pigs containing soft tissues (R2 = 0.98) without requiring knowledge of true density or phantom calibration of the scans. DISCUSSION: BRP allows automated comparison of image data from different image modalities (pQCT, µCT) using different energy settings, in archeological bone and wet specimens. The method enables low-resolution data generated in vivo to be compared with the fossil and archaeological record. Such experimental approaches would substantially improve behavioral inferences based on trabecular bone microstructure.

StW 573 Australopithecus prometheus: Its Significance for an Australopith Bauplan.

The StW 573 skeleton of Australopithecus prometheus from Sterkfontein Member 2 is some 93% complete and thus by far the most complete member of that genus yet found. Firmly dated at 3.67 Ma, it is one of the earliest specimens of its genus. A crucial aspect of interpretation of locomotor behaviour from fossil remains is an understanding of the palaeoenvironment in which the individual lived and the manner in which it would have used it. While the value of this ecomorphological approach is largely accepted, it has not been widely used as a stable framework on which to build evolutionary biomechanical interpretations. Here, we collate the available evidence on StW 573's anatomy in order, as far as currently possible, to reconstruct what might have been this individual's realized and potential niche. We explore the concept of a common Australopithecus "bauplan" by comparing the morphology and ecological context of StW 573 to that of paenocontemporaneous australopiths including Australopithecus anamensis and KSD-VP-1/1 Australopithecus afarensis. Each was probably substantially arboreal and woodland-dwelling, relying substantially on arboreal resources. We use a hypothesis-driven approach, tested by: virtual experiments, in the case of extinct species; biomechanical analyses of the locomotor behaviour of living great ape species; and analogical experiments with human subjects. From these, we conclude that the habitual locomotor mode of all australopiths was upright bipedalism, whether on the ground or on branches. Some later australopiths such as Australopithecus sediba undoubtedly became more terrestrial, allowing sacrifice of arboreal stability in favour of manual dexterity. Indeed, modern humans retain arboreal climbing skills but have further sacrificed arboreal effectiveness for enhanced ability to sustain striding terrestrial bipedalism over much greater distances. We compare StW 573's locomotor adaptations to those of living great apes and protohominins, and agree with those earlier observers who suggest that the common panin-hominin last common ancestor was postcranially more like Gorilla than Pan.

Morphometric analysis of the hominin talus: Evolutionary and functional implications.

The adoption of bipedalism is a key benchmark in human evolution that has impacted talar morphology. Here, we investigate talar morphological variability in extinct and extant hominins using a 3D geometric morphometric approach. The evolutionary timing and appearance of modern human-like features and their contributions to bipedal locomotion were evaluated on the talus as a whole, each articular facet separately, and multiple combinations of facets. Distinctive suites of features are consistently present in all fossil hominins, despite the presence of substantial interspecific variation, suggesting a potential connection of these suites to bipedal gait. A modern human-like condition evolved in navicular and lateral malleolar facets early in the hominin lineage compared with other facets, which demonstrate more complex morphological variation within Homininae. Interestingly, navicular facet morphology of Australopithecus afarensis is derived in the direction of Homo, whereas more recent hominin species such as Australopithecus africanus and Australopithecus sediba retain more primitive states in this facet. Combining the navicular facet with the trochlea and the posterior calcaneal facet as a functional suite, however, distinguishes Australopithecus from Homo in that the medial longitudinal arch had not fully developed in the former. Our results suggest that a more everted foot and stiffer medial midtarsal region are adaptations that coincide with the emergence of bipedalism, whereas a high medial longitudinal arch emerges later in time, within Homo. This study provides novel insights into the emergence of talar morphological traits linked to bipedalism and its transition from a facultative to an obligate condition.

An examination of the cross-sectional geometrical properties of the long bone diaphyses of Holocene foragers from Roonka, South Australia.

OBJECTIVES: This study examines long bone diaphyseal rigidity and shape of hunter-gatherers at Roonka to make inferences about subsistence strategies and mobility of inhabitants of semi-arid southeastern Australia. Roonka is a cemetery site adjacent to the Lower Murray River, which contains over 200 individuals buried throughout the Holocene. Archaeological evidence indicates that populations living near this river corridor employed mobile, risk averse foraging strategies. METHODS: This prediction of lifestyle was tested by comparing the cross-sectional geometric properties of the humerus, radius, ulna, femur, tibia, and fibula of individuals from Roonka to samples of varying subsistence strategies. Bilateral asymmetry of the upper limb bones was also examined. RESULTS: Roonka males and females have moderately high lower limb diaphyseal rigidity and shape. In the upper limb, females have low rigidity and bilateral asymmetry while males have moderately high rigidity and bilateral asymmetry. This pattern is similar to other foraging groups from Australia and southern Africa that have behaviorally adapted to arid and semi-arid environments. DISCUSSION: Lower limb results suggest that populations in the Lower Murray River Valley had relatively elevated foraging mobility. Upper limb rigidity and bilateral asymmetry indicate a sexual division of labor at Roonka. Females resemble other samples that had mixed subsistence strategies that involved hunting, gathering, and processing tasks. Males display a pattern similar to groups that preferentially hunted large game, but that supplemented this source with smaller game and riverine resources.

The upper limb skeleton and behavioral lateralization of modern humans from Zhaoguo Cave, southwestern China.

OBJECTIVES: Aims of the study are to initially describe and comparatively evaluate the morphology of the new Zhaoguo M1 upper limb remains, and contextualize upper limb functional adaptations among those of other worldwide Upper Paleolithic (UP) humans to make inferences about subsistence-related activity patterns in southwestern China at the Pleistocene-Holocene boundary. MATERIALS AND METHODS: The preserved Zhaoguo M1 skeletal remains include paired humeri, ulnae, and radii, among others. These specimens were scanned using micro-computed tomography to evaluate internal structural properties, while external osteometric dimensions of the Zhaoguo M1 upper limb elements also were acquired. Both sets of measurements were compared to published data on Neandertals, and Middle and Upper Paleolithic modern humans. RESULTS: The upper limb elements of Zhaoguo M1 display a suite of characteristics that generally resemble those of other contemporary Late UP (LUP) modern humans, while robusticity indices generally fall within the upper range of LUP variation. The Zhaoguo M1 upper limb elements display fewer traits resembling those of late archaic humans. The Zhaoguo M1 individual exhibits diaphyseal asymmetry in several upper limb elements suggesting left hand dominance. When evaluating the full range of magnitudes of humeral bilateral asymmetry in the comparative sample, Zhaoguo M1 falls at the lower end overall, but yet is relatively higher than contemporary LUP modern humans specifically from East Eurasia. DISCUSSION: The Zhaoguo M1 individual suggests typical LUP modern human upper limb morphology persisted in southwest China until the end of the last glacial period. Upper limb bone asymmetry of Zhaoguo M1 also indicates that behavioral activities attributed to a hunter-gatherer tradition apparently extended through the Pleistocene-Holocene transition in this region.

The influence of mobility strategy on the modern human talus.

OBJECTIVES: The primate talus is known to have a shape that varies according to differences in locomotion and substrate use. While the modern human talus is morphologically specialized for bipedal walking, relatively little is known on how its morphology varies in relation to cultural and environmental differences across time. Here we compare tali of modern human populations with different subsistence economies and lifestyles to explore how cultural practices and environmental factors influence external talar shape. MATERIALS AND METHODS: The sample consists of digital models of 142 tali from 11 archaeological and post-industrial modern human groups. Talar morphology was investigated through 3D (semi)landmark based geometric morphometric methods. RESULTS: Our results show distinct differences between highly mobile hunter-gatherers and more sedentary groups belonging to a mixed post-agricultural/industrial background. Hunter-gatherers exhibit a more "flexible" talar shape, everted posture, and a more robust and medially oriented talar neck/head, which we interpret as reflecting long-distance walking strictly performed barefoot, or wearing minimalistic footwear, along uneven ground. The talus of the post-industrial population exhibits a "stable" profile, neutral posture, and a less robust and orthogonally oriented talar neck/head, which we interpret as a consequence of sedentary lifestyle and use of stiff footwear. DISCUSSION: We suggest that talar morphological variation is related to the adoption of constraining footwear in post-industrial society, which reduces ankle range of motion. This contrasts with hunter-gatherers, where talar shape shows a more flexible profile, likely resulting from a lack of footwear while traversing uneven terrain. We conclude that modern human tali vary with differences in locomotor and cultural behavior.

The long limb bones of the StW 573 Australopithecus skeleton from Sterkfontein Member 2: Descriptions and proportions.

Due to its completeness, the A.L. 288-1 ('Lucy') skeleton has long served as the archetypal bipedal Australopithecus. However, there remains considerable debate about its limb proportions. There are three competing, but not necessarily mutually exclusive, explanations for the high humerofemoral index of A.L. 288-1: (1) a retention of proportions from an Ardipithecus-like chimp/human last common ancestor (CLCA); (2) indication of some degree of climbing ability; (3) allometry. Recent discoveries of other partial skeletons of Australopithecus, such as those of Australopithecus sediba (MH1 and MH2) and Australopithecus afarensis (KSD-VP-1/1 and DIK-1/1), have provided new opportunities to test hypotheses of early hominin body size and limb proportions. Yet, no early hominin is as complete (>90%), as is the ∼3.67 Ma 'Little Foot' (StW 573) skeleton from Sterkfontein Member 2. Here, we provide the first descriptions of its upper and lower long limb bones, as well as a comparative context of its limb proportions. We found that StW 573 possesses absolutely longer limb lengths than A.L. 288-1, but both skeletons show similar limb proportions. This finding seems to argue against a purely allometric explanation for A.L. 288-1 limb proportions. In fact, our multivariate allometric analysis suggests that limb lengths of Australopithecus, as represented by StW 573 and A.L. 288-1, exhibit a significantly different (p < 0.001) allometric pattern than that which typifies modern humans and African apes. Like some previous analyses, our results also suggest that hominin limb evolution occurred in two stages with: first, a modest increase in lower limb length and a concurrent shortening of the antebrachium between Ardipithecus and Australopithecus, followed by a considerable lengthening of the lower limb along with a decrease of both upper limb elements occurring between Australopithecus and Homo sapiens.

The bony labyrinth of StW 573 ("Little Foot"): Implications for early hominin evolution and paleobiology.

Because of its exceptional degree of preservation and its geological age of ∼3.67 Ma, StW 573 makes an invaluable contribution to our understanding of early hominin evolution and paleobiology. The morphology of the bony labyrinth has the potential to provide information about extinct primate taxonomic diversity, phylogenetic relationships and locomotor behaviour. In this context, we virtually reconstruct and comparatively assess the bony labyrinth morphology in StW 573. As comparative material, we investigate 17 southern African hominin specimens from Sterkfontein, Swartkrans and Makapansgat (plus published data from two specimens from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant chimpanzee specimens. We apply a landmark-based geometric morphometric method for quantitatively assessing labyrinthine morphology. Morphology of the inner ear in StW 573 most closely resembles that of another Australopithecus individual from Sterkfontein, StW 578, recovered from the Jacovec Cavern. Within the limits of our sample, we observe a certain degree of morphological variation in the Australopithecus assemblage of Sterkfontein Member 4. Cochlear morphology in StW 573 is similar to that of other Australopithecus as well as to Paranthropus specimens included in this study, but it is substantially different from early Homo. Interestingly, the configuration of semicircular canals in Paranthropus specimens from Swartkrans differs from other fossil hominins, including StW 573. Given the role of the cochlea in the sensory-driven interactions with the surrounding environment, our results offer new perspectives for interpreting early hominin behaviour and ecology. Finally, our study provides additional evidence for discussing the phylogenetic polarity of labyrinthine traits in southern African hominins.

The endocast of StW 573 ("Little Foot") and hominin brain evolution.

One of the most crucial debates in human paleoneurology concerns the timing and mode of the emergence of the derived cerebral features in the hominin fossil record. Given its exceptional degree of preservation and geological age (i.e., 3.67 Ma), StW 573 ('Little Foot') has the potential to shed new light on hominin brain evolution. Here we present the first detailed comparative description of the external neuroanatomy of StW 573. The endocast was virtually reconstructed and compared to ten southern African hominin specimens from Makapansgat, Malapa, Sterkfontein and Swartkrans attributed to Australopithecus and Paranthropus. We apply an automatic method for the detection of sulcal and vascular imprints. The endocranial surface of StW 573 is crushed and plastically deformed in a number of locations. The uncorrected and therefore minimum cranial capacity estimate is 408 cm3 and plots at the lower end of Australopithecus variation. The endocast of StW 573 approximates the rostrocaudally elongated and dorsoventrally flattened endocranial shape seen in Australopithecus and displays a distinct left occipital petalia. StW 573 and the comparative early hominin specimens share a similar sulcal pattern in the inferior region of the frontal lobes that also resembles the pattern observed in extant chimpanzees. The presumed lunate sulcus in StW 573 is located above the sigmoid sinus, as in extant chimpanzees, while it is more caudally positioned in SK 1585 and StW 505. The middle branch of the middle meningeal vessels derives from the anterior branch, as in MH 1, MLD 37/38, StW 578. Overall, the cortical anatomy of StW 573 displays a less derived condition compared to the late Pliocene/early Pleistocene southern African hominins (e.g., StW 505, SK 1585).

Complex variation of trabecular bone structure in the proximal humerus and femur of five modern human populations.

OBJECTIVE: This project investigates trabecular bone structural variation in the proximal humerus and femur of hunter-gatherer, mixed-strategy agricultural, medieval, and human groups to address three questions: (a) What is the extent of trabecular bone structural variation in the humerus and femur between populations with different inferred activity levels? (b) How does variation in the proximal humerus relate to variation in the proximal femur? (c) Are trabecular bone microstructural variables sexually dimorphic? METHODS: The proximal humerus and femur of 73 adults from five human groups with distinct subsistence strategies were scanned using a micro-computed tomography system. Centralized volumes of interest within the humeral and femoral heads were extracted and analyzed to quantify bone volume fraction, trabecular thickness, trabecular separation, connectivity density, degree of anisotropy, and bone surface density. RESULTS: In the humerus and femur, groups with the highest inferred activity levels have higher bone volume fraction and trabecular thickness, and lower bone surface density than those with lower inferred activity levels. However, the humeral pattern does not exactly mirror that of the femur, which demonstrates a steeper gradient of difference between subsistence groups. No significant differences were identified in trabecular separation. No consistent patterns of sexual dimorphism were present in the humerus or femur. CONCLUSIONS: Reduced skeletal robusticity of proximal humeral and femoral trabecular bone corresponds with reduced activity level inferred from subsistence strategy. However, human trabecular bone structural variation is complex and future work should explore how other factors (diet, climate, genetics, disease load, etc.), in addition to activity, influence bone structural variation.

Human-like hip joint loading in Australopithecus africanus and Paranthropus robustus.

Adaptations indicative of habitual bipedalism are present in the earliest recognized hominins. However, debate persists about various aspects of bipedal locomotor behavior in fossil hominins, including the nature of gait kinematics, locomotor variability across different species, and the degree to which various australopith species engaged in arboreal behaviors. In this study, we analyze variation in trabecular bone structure of the femoral head using a sample of modern humans, extant non-human hominoids, baboons, and fossil hominins attributed to Australopithecus africanus, Paranthropus robustus, and the genus Homo. We use µCT data to characterize the fabric anisotropy, material orientation, and bone volume fraction of trabecular bone to reconstruct hip joint loading conditions in these fossil hominins. Femoral head trabecular bone fabric structure in australopiths is more similar to that of modern humans and Pleistocene Homo than extant apes, indicating that these australopith individuals walked with human-like hip kinematics, including a more limited range of habitual hip joint postures (e.g., a more extended hip) during bipedalism. Our results also indicate that australopiths have robust femoral head trabecular bone, suggesting overall increased loading of the musculoskeletal system comparable to that imposed by extant apes. These results provide new evidence of human-like bipedal locomotion in Pliocene hominins, even while other aspects of their musculoskeletal systems retain ape-like characteristics.

Cranial vault thickness variation and inner structural organization in the StW 578 hominin cranium from Jacovec Cavern, South Africa.

The Sterkfontein Caves site is one of the richest early hominin fossil localities in Africa. More specifically, the fossiliferous deposits within the lower-lying Jacovec Cavern have yielded valuable hominin remains; prominent among them is the Australopithecus partial cranium StW 578. Due to the fragmentary nature of the braincase, the specimen has not yet been formally assigned to a species. In this context, we employ microtomography to quantify cranial thickness and composition of StW 578 in order to assess its taxonomic affinity. As comparative material, we investigate 10 South African hominin cranial specimens from Sterkfontein (StW 505, Sts 5, Sts 25, Sts 71), Swartkrans (SK 46, SK 48, SK 49) and Makapansgat (MLD 1, MLD 10, MLD 37/38), attributed to either Australopithecus or Paranthropus, as well as 10 extant human and 10 extant chimpanzee crania. Thickness variation in and structural arrangement of the inner and outer cortical tables and the diploë are automatically assessed at regular intervals along one parasagittal and one coronal section. Additionally, topographic cranial vault thickness distribution is visualized using color maps. Comparisons highlight an absolutely and relatively thickened condition of the StW 578 cranial vault versus those of other South African Plio-Pleistocene hominins. Moreover, in StW 578, as well as in the Australopithecus specimens Sts 5 and Sts 71 from Sterkfontein, the diploic layer contributes substantially to cumulative vault thickness (i.e., >60%). Within the comparative sample investigated here, StW 505 and Sts 71 from Sterkfontein Member 4, both attributed to Australopithecus, most closely resemble StW 578 in terms of cranial vault thickness values, tissue proportions, and two- and three-dimensional distributions. Including additional Plio-Pleistocene Australopithecus and Paranthropus crania from South and East Africa in future studies would further help establish morphological variability in these hominin taxa.

Inter-ray variation in metatarsal strength properties in humans and African apes: Implications for inferring bipedal biomechanics in the Olduvai Hominid 8 foot.

When measured as a ratio of mean midshaft diameter to bone length, the OH 8 fossil hominin foot exhibits a metatarsal (Mt) robusticity pattern of 1 > 5 > 3 > 4 > 2, which differs from the widely perceived "common" modern human pattern (1 > 5 > 4 > 3 > 2); African apes generally exhibit a third pattern (1 > 2 > 3 > 4 > 5). Largely because of the relative ranking of Mt2 and Mt5, OH 8 metatarsals structurally resemble the pattern exhibited by bipedal humans more than the pattern of quadrupedal and climbing African apes. Considering only these three phenotypes, however, discounts the potentially important functional implications of variation in modern human (and African ape) metatarsal robusticity patterns, suggesting that they are not useful for interpreting the specific biomechanics of a bipedal gait in fossils (i.e., whether it was modern human-like or not). Using computed tomography scans to quantify metatarsal midshaft cross-sectional geometry in a large sample of Homo (n=130), Gorilla (n=44) and Pan (n=80), we documented greater variation in metatarsal robusticity patterns than previously recognized in all three groups. While apes consistently show a 1 > 2 > 3 > 4 > 5 pattern in our larger sample, there does not appear to be a similarly precise single "common" human pattern. Rather, human metatarsals converge towards a 1 > 4/5 > 2/3 pattern, where metatarsals 4 and 5, and metatarsals 2 and 3, often "flip" positions relative to each other depending on the variable examined. After reassessing what a "common" human pattern could be based on a larger sample, the previously described OH 8 pattern of 1 > 5 > 3 > 4 > 2 is only observed in some humans (<6%) and almost never in apes (<0.5%). Although this suggests an overall greater similarity to (some) humans than to any ape in loading of the foot, the relatively rare frequency of these humans in our sample underscores potential differences in loading experienced by the medial and lateral columns of the OH 8 foot compared to modern humans.