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Competing Interests: Declaration of competing interest The authors declare no competing interests.

Reconstruction of the locomotor repertoire of the australopiths (Australopithecus and Paranthropus) has progressively integrated information from the mechanosensitive internal structure of the appendicular skeleton. Recent investigations showed that the arrangement of the trabecular network at the femoral head center is biomechanically compatible with the pattern of cortical bone distribution across the neck, both suggesting a full commitment to bipedalism in australopiths, but associated with a slightly altered gait kinematics compared to Homo involving more lateral deviation of the body center of mass over the stance limb. To provide a global picture in Paranthropus robustus of the trabecular architecture of the proximal femur across the head, neck and greater trochanter compartments, we applied techniques of virtual imaging to the variably preserved Early Pleistocene specimens SK 82, SK 97, SK 3121, SKW 19 and SWT1/LB-2 from the cave site of Swartkrans, South Africa. We also assessed the coherence between the structural signals from the center of the head and those from the trabecular network of the inferolateral portion of the head and the inferior margin of the neck, sampling the so-called vertical bundle, which in humans represents the principal compressive system of the joint. Our analyses show a functionally related trabecular organization in Pa. robustus that closely resembles the extant human condition, but which also includes some specificities in local textural arrangement. The network of the inferolateral portion of the head shows a humanlike degree of anisotropy and a bone volume fraction intermediate between the extant human and the African ape patterns. These results suggest slight differences in gait kinematics between Pa. robustus and extant humans. The neck portion of the vertical bundle revealed a less biomechanically sensitive signal. Future investigations on the australopith hip joint loading environment should more carefully investigate the trabecular structure of the trochanteric region and possible structural covariation between cortical bone distribution across the neck and site-specific trabecular properties of the arcuate bundle., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

A partial left femur (TM 266-01-063) was recovered in July 2001 at Toros-Menalla, Chad, at the same fossiliferous location as the late Miocene holotype of Sahelanthropus tchadensis (the cranium TM 266-01-060-1). It was recognized as a probable primate femur in 2004 when one of the authors was undertaking a taphonomic survey of the fossil assemblages from Toros-Menalla. We are confident the TM 266 femoral shaft belongs to a hominid. It could sample a hominid hitherto unrepresented at Toros-Menalla, but a more parsimonious working hypothesis is that it belongs to S. tchadensis. The differences between TM 266 and the late Miocene Orrorin tugenensis partial femur BAR 1002'00, from Kenya, are consistent with maintaining at least a species-level distinction between S. tchadensis and O. tugenensis. The results of our preliminary functional analysis suggest the TM 266 femoral shaft belongs to an individual that was not habitually bipedal, something that should be taken into account when considering the relationships of S. tchadensis. The circumstances of its discovery should encourage researchers to check to see whether there is more postcranial evidence of S. tchadensis among the fossils recovered from Toros-Menalla., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Studies of the australopith (Australopithecus and Paranthropus) proximal femur have increasingly integrated information from the local arrangement of the cortical and cancellous bone to allow functional-biomechanical inferences on the locomotor behavioral patterns. In Australopithecus africanus and Paranthropus robustus, the cancellous bone organization at the center of the femoral head shows principal strut orientation similar to that of fossil and recent humans, which indicates that australopiths were human-like in many aspects of their bipedalism. However, by combining outer morphology with superoinferior asymmetry in cortical bone thickness at the base of neck and mid-neck, it has been suggested that, while adapted for terrestrial bipedality, australopiths displayed a slightly altered gait kinematics compared to Homo. We used techniques of 2D and 3D virtual imaging applied to an X-ray microtomographic record to assess cortical bone distribution along the entire femoral neck compartment in four upper femora from Swartkrans, South Africa (SK 82, SK 97, SK 3121, and SWT1/LB-2) and compared the results to the extant human and chimpanzee conditions. Our results support and extend previous evidence for more symmetric superior and inferior femoral neck cortical thicknesses in P. robustus than in modern humans and show that the differences are even greater than previously reported. However, P. robustus and humans still share a trend of lateral-to-medial decrease in asymmetry of the superior/inferior cortical thickness ratio, while this pattern is reversed in chimpanzees. We also identified two features uniquely characterizing P. robustus: an accentuated contrast between the relatively thicker anterior and the thinner posterior walls, and a more marked lateral-to-medial thinning of both cortices compared to extant humans and chimpanzees, which indicate wider interspecific differences among hominids in structural organization of the proximal femur than previously reported. It remains to be ascertained if, and to what extent, these features also characterize the femoral neck of Australopithecus., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Wezmeh Cave, in the Kermanshah region of Central Western Zagros, Iran, produced a Late Pleistocene faunal assemblage rich in carnivorans along with a human right maxillary premolar, Wezmeh 1, an unerupted tooth from an 8 ± 2 year-old individual. Uranium-series analyses of the fauna by alpha spectrometry provided age estimates between 70 and 11 ka. Crown dimensions place the tooth specimen at the upper limits of Late Pleistocene human ranges of variation. Wezmeh 1 metameric position (most likely a P 3 ) remains uncertain and only its surficial morphology has been described so far. Accordingly, we used microfocus X-ray tomography (12.5 μm isotropic voxel size) to reassess the metameric position and taxonomic attribution of this specimen. We investigated its endostructural features and quantified crown tissue proportions. Topographic maps of enamel thickness (ET) distribution were also generated, and semilandmark-based geometric morphometric analyses of the enamel-dentine junction (EDJ) were performed. We compared Wezmeh 1 with unworn/slightly-moderately worn P 3 and P 4 of European Neanderthals, Middle Paleolithic modern humans from Qafzeh, an Upper Paleolithic premolar, and Holocene humans. The results confirm that Wezmeh 1 represents a P 3 . Based on its internal conformation and especially EDJ shape, Wezmeh 1 aligns closely with Neanderthals and is distinct from the fossil and extant modern human pattern of our comparative samples. Wezmeh 1 is thus the first direct evidence of Neanderthal presence on the western margin of the Iranian Plateau., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Tooth wear is an important feature for reconstructing diet, food processing and cultural habits of past human populations. In particular, occlusal wear facets can be extremely useful for detecting information about diet and non-masticatory behaviors. The aim of this study is to reconstruct the diet and cultural behavior of the Neanderthal specimen Regourdou 1 (Dordogne, Southern France) from the analysis of the macrowear pattern, using the occlusal fingerprint analysis method. In addition, we have also examined whether there is any association between the observed dental macrowear and mandibular bone distribution and root dentine thickness. The posterior dentition of Regourdou 1 is characterized by an asymmetric wear pattern, with the right side significantly more worn than the left. In contrast, the left lower P 3 shows a more advanced wear than the right premolar, with unusual semicircular enamel wear facets. The results from occlusal fingerprint analysis of this unique pattern suggest tooth-tool uses for daily task activities. Moreover, the left buccal aspect of the mandibular cortical bone is thicker than its right counterpart, and the left P 3 has a thicker radicular dentine layer than its antimere. These results show a certain degree of asymmetry in cortical bone topography and dentine tissue that could be associated with the observed dental macrowear pattern. The molar macrowear pattern also suggests that Regourdou 1 had a mixed diet typical of those populations living in temperate deciduous woodlands and Mediterranean habitats, including animal and plant foods. Although this study is limited to one Neanderthal individual, future analyses based on a larger sample may further assist us to better understand the existing relationship between mandibular architecture, occlusal wear and the masticatory apparatus in humans., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Here we analyze 1.07-0.99 million-year-old pelvic remains UA 173/405 from Buia, Eritrea. Based on size metrics, UA 173/405 is likely associated with an already described pubic symphysis (UA 466) found nearby. The morphology of UA 173/405 was quantitatively characterized using three-dimensional landmark-based morphometrics and linear data. The Buia specimen falls within the range of variation of modern humans for all metrics investigated, making it unlikely that the shared last common ancestor of Late Pleistocene Homo species would have had an australopith-like pelvis. The discovery of UA 173/405 adds to the increasing number of fossils suggesting that the postcranial morphology of Homo erectus s.l. was variable and, in some cases, nearly indistinguishable from modern human morphology. This Eritrean fossil demonstrates that modern human-like pelvic morphology may have had origins in the Early Pleistocene, potentially within later African H. erectus., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Locality 1, in the Lower Cave of the Zhoukoudian cave complex, China, is one of the most important Middle Pleistocene paleoanthropological and archaeological sites worldwide, with the remains of c. 45 Homo erectus individuals, 98 mammalian taxa, and thousands of lithic tools recovered. Most of the material collected before World War II was lost. However, besides two postcranial elements rediscovered in China in 1951, four human permanent teeth from the 'Dragon Bone Hill,' collected by O. Zdansky between 1921 and 1923, were at the time brought to the Paleontological Institute of Uppsala University, Sweden, where they are still stored. This small sample consists of an upper canine (PMU 25719), an upper third molar (PMU M3550), a lower third premolar crown (PMU M3549), and a lower fourth premolar (PMU M3887). Some researchers have noted the existence of morpho-dimensional differences between the Zhoukoudian and the H. erectus dental assemblage from Sangiran, Java. However, compared to its chrono-geographical distribution, the Early to Middle Pleistocene dental material currently forming the Chinese-Indonesian H. erectus hypodigm is quantitatively meager and still poorly characterized for the extent of its endostructural variation. We used micro-focus X-ray tomography techniques of virtual imaging coupled with geometric morphometrics for comparatively investigating the endostructural conformation (tissue proportions, enamel thickness distribution, enamel-dentine junction morphology, pulp cavity shape) of the four specimens stored in Uppsala, all previously reported for their outer features. The results suggest the existence of time-related differences between continental and insular Southeast Asian dental assemblages, the Middle Pleistocene Chinese teeth apparently retaining an inner signature closer to the likely primitive condition represented by the Early Pleistocene remains from Java, while the Indonesian stock evolved toward tooth structural simplification., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The original hominin femur (Femur I) and calotte discovered at Trinil, Java by Eugene Dubois in 1891/1892 played a key role in the early history of human paleontology by purportedly demonstrating the contemporaneity of archaic cranial form with modern human erect (bipedal) posture. On this basis, both specimens were subsequently assigned to Pithecanthropus erectus, later transferred to Homo erectus. However, chronological and phylogenetic links between the two have been questioned from the beginning. Four additional hominin partial femora (Femora II-V) from Trinil were subsequently described but have played a relatively minor part in evolutionary scenarios. Here we present the results of a new analysis of structural and density characteristics of the Trinil femora obtained using computed tomography. Trinil Femur I shows none of the characteristics typical of early Homo femora from elsewhere in Asia or Africa, including a relatively long neck, increased mediolateral bending rigidity of the mid-proximal shaft, or a low position of minimum mediolateral breath on the shaft. In contrast, Femora II-V all demonstrate features that are more consistent with this pattern. In addition, material density distributions within the specimens imply more recent and less complete fossilization of Femur I than Femora II-V. Thus, it is very likely that Trinil Femur I derives from a much more recent time period than the calotte, while the less famous and less complete Femora II-V may represent H. erectus at Trinil. The morphological variation within the Trinil femora can be attributed to broader changes in pelvic morphology occurring within the Homo lineage between the Early and late Middle Pleistocene., (Copyright © 2015. Published by Elsevier Ltd.)

Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has significantly enlarged the fossil record of Homo erectus sensu lato (s.l.). Additional evidence comes from the Danakil Depression of Eritrea, where over 200 late Early to early Middle Pleistocene sites have been identified within a ∼1000 m-thick sedimentary succession outcropping in the Dandiero Rift Basin, near Buia. Along with an adult cranium (UA 31), which displays a blend of H. erectus-like and derived morpho-architectural features and three pelvic remains, two isolated permanent incisors (UA 222 and UA 369) have also been recovered from the 1 Ma (millions of years ago) Homo-bearing outcrop of Uadi Aalad. Since 2010, our surveys have expanded to the nearby (4.7 km) site of Mulhuli-Amo (MA). This is a fossiliferous area that has been preliminarily surveyed because of its exceptional concentration of Acheulean stone tools. So far, the site has yielded 10 human remains, including the unworn crown of a lower permanent molar (MA 93). Using diverse analytical tools (including high resolution μCT and μMRI), we analysed the external and internal macromorphology and microstructure of the three specimens, and whenever possible compared the results with similar evidence from early Homo, H. erectus s.l., H. antecessor, H. heidelbergensis (from North Africa), Neanderthals and modern humans. We also assessed the UA 369 lower incisor from Uadi Aalad for root completion timing and showed that it compares well with data for root apex closure in modern human populations., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The biomechanical characterization of lower limb long bones in the chrono-ecogeographically diverse species Homo erectus is a fundamental step for assessing evolutionary changes in locomotor mode and body shape that occurred within the genus Homo. However, the samples available for the Early and earlier Middle Pleistocene are small and widely scattered in time and space, thus limiting our understanding of the nature and polarity of morphological trends. Compared to the African fossil record, loading histories based on detailed biomechanical assessment of diaphyseal strength in Indonesian H. erectus lower limb long bones have not been assessed. By using a microtomographic record (μCT), we performed a quantitative analysis of the biomechanical properties and structural organization of Kresna 11, a late Early Pleistocene adult H. erectus femoral shaft from the Sangiran Dome, Central Java. Relative to the modern human condition, Kresna 11 shows the predominant mediolateral cortical thickening (hypertrophy) and the distal displacement of the minimum diaphyseal breadth characteristic of early Homo femora, associated nonetheless with relatively modest cortical thickness within the mid-proximal portion. Synthetic functional imaging of the shaft through the planar representation of its inner structure has revealed distal thickening of the medial cortex, a feature previously unreported in H. erectus. The increase in relative mediolateral bending strength observed in Kresna 11 supports the hypothesis that, rather than simply reflecting differences in patterns of locomotor loading, biomechanical properties of the femoral shaft in archaic Homo are strongly influenced by body shape, i.e., variations in pelvic breadth and femoral neck length., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The recovery at Shi'bat Dihya 1 (SD1) of a dense Middle Paleolithic human occupation dated to 55 ka BP sheds new light on the role of the Arabian Peninsula at the time of the alleged expansion of modern humans out of Africa. SD1 is part of a complex of Middle Paleolithic sites cut by the Wadi Surdud and interstratified within an alluvial sedimentary basin in the foothills that connect the Yemeni highlands with the Tihama coastal plain. A number of environmental proxies indicate arid conditions throughout a sequence that extends between 63 and 42 ka BP. The lithic industry is geared toward the production of a variety of end products: blades, pointed blades, pointed flakes and Levallois-like flakes with long unmodified cutting edges, made from locally available rhyolite. The occasional exploitation of other local raw materials, that fulfill distinct complementary needs, highlights the multi-functional nature of the occupation. The slightly younger Shi'bat Dihya 2 (SD2) site is characterized by a less elaborate production of flakes, together with some elements (blades and pointed flakes) similar to those found at SD1, and may indicate a cultural continuity between the two sites. The technological behaviors of the SD1 toolmakers present similarities with those documented from a number of nearly contemporaneous assemblages from southern Arabia, the Levant, the Horn of Africa and North Africa. However, they do not directly conform to any of the techno-complexes typical of the late Middle Paleolithic or late Middle Stone Age from these regions. This period would have witnessed the development of local Middle Paleolithic traditions in the Arabian Peninsula, which suggests more complex settlement dynamics and possible population interactions than commonly inferred by the current models of modern human expansion out of Africa., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The assessment of the degree of similarity or difference between Neanderthals and modern humans in their patterns of dental development remains a controversial matter. Here we report results from the microtomographic-based (SR-microCT) high-resolution structural investigation of the maxilla and mandible of the Neanderthal child from Roc de Marsal, Dordogne, France (likely from OIS 5a). Following their virtual extraction and 3D rendering, we assessed the maturational stage of each of the 41 dental elements (20 deciduous and 21 permanent) forming its mixed dentition. By using a Bayesian approach, we calculated the probability that its deciduous and permanent mandibular sequences are found within the extant human variation as illustrated by a tomographic CT-based sub-sample of 32 children (deciduous dentition) and a panoramic radiographic- and CT-based whole sample of 343 living children (permanent dentition). Results show that neither the deciduous nor the permanent mandibular sequences displayed by Roc de Marsal are precisely found within our modern comparative files. In both sequences, the most influential factor is represented by a slight discrepancy in the Neanderthal child between the stage of mineralization of the first molar, which is proportionally advanced, and the maturational level reached by its incisors, which are proportionally delayed. Following a quantitative volumetric analysis of the deciduous teeth, we suggest that this characteristic may be related to differences between Neanderthals and modern humans in absolute dental size and relative size proportions between front and cheek teeth, as well as to structural differences in dental tissue proportions.

The thickness of dental enamel is often discussed in paleoanthropological literature, particularly with regard to differences in growth, health, and diet between Neandertals and modern humans. Paleoanthropologists employ enamel thickness in paleodietary and taxonomic studies regarding earlier hominins, but variation in enamel thickness within the genus Homo has not been thoroughly explored despite its potential to discriminate species and its relevance to studies of growth and development. Radiographic two-dimensional studies indicate that Neandertal molar enamel is thin relative to the thick enamel of modern humans, although such methods have limited accuracy. Here we show that, measured via accurate high-resolution microtomographic imaging, Neandertal molar enamel is absolutely and relatively thinner than modern human enamel at most molar positions. However, this difference relates to the ratio of coronal dentine volume to total crown volume, rather than the quantity of enamel per se. The absolute volume of Neandertal molar enamel is similar to that of modern humans, but Neandertal enamel is deposited over a larger volume of coronal dentine, resulting in lower average (and relative) enamel thickness values. Sample sizes do not permit rigorous intragroup comparisons, but Neandertal molar tissue proportions evince less variation than the modern human sample. Differences in three- and two-dimensional enamel thickness data describing Neandertal molars may be explained by dimensional reduction. Although molar tissue proportions distinguish Neanderthals from recent Homo sapiens, additional study is necessary to assess trends in tissue proportions in the genus Homo throughout the Pleistocene.

Cancellous bone retains structural and behavioural properties which are time and strain-rate dependent. As the orientation of the trabeculae (trajectories) follows the direction of the principal strains imposed by daily loadings, habitual postural and locomotor behaviours are responsible for a variety of trabecular architectures and site-specific textural arrangements of the pelvic cancellous network. With respect to the great ape condition, the human trabecular pattern is characterized by a distinctive ilioischial bundle, an undivided sacropubic bundle, and a full diagonal crossing (approximately 100 degrees) over the acetabulum between the ilioischial and the sacropubic bundles. Advanced digital image processing (DIP) of hip bone radiographs has revealed that adolescent and adult South African australopithecines retained an incompletely developed human-like trabecular pattern associated with gait-related features that are unique among the extant primates.