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The postcranial skeleton of Australopithecus afarensis (AL 288-1) exhibits clear adaptations for bipedality, although there is some debate as to the efficiency and frequency of such upright movement. Some researchers argue that AL 288-1 walked with an erect limb like modern humans do, whilst others advocate for a "bent-hip bent-knee" (BHBK) gait, although in recent years the general consensus favors erect bipedalism. To date, no quantitative method has addressed the articulation of the AL 288-1 hip joint, nor its range of motion (ROM) with consideration for joint spacing, used as a proxy for the thickness of the articular cartilage present within the joint spacing which can affect how a joint moves. Here, we employed ROM mapping methods to estimate the joint spacing of AL 288-1's hip joint in comparison to a modern human and chimpanzee. Nine simulations assessed different joint spacing and tested the range of joint congruency (i.e., ranging from a closely packed socket to loosely packed). We further evaluated the sphericity of the femoral head and whether three rotational degrees of freedom (DOFs) sufficiently captures the full ROM or if translational DOFs must be included. With both setups, we found that the AL 288-1 hip was unlikely to be highly congruent (as it is in modern humans) because this would severely restrict hip rotational movement and would severely limit the capability for both bipedality and even arboreal locomotion. Rather, the hip was more cartilaginous than it is in the modern humans, permitting the hip to rotate into positions necessitated by both terrestrial and arboreal movements. Rotational-only simulations found that AL 288-1 was unable to extend the hip like modern humans, forcing the specimen to employ a BHBK style of walking, thus contradicting 40+ years of previous research into the locomotory capabilities of AL 288-1. Therefore, we advocate that differences in the sphericity of the AL 288-1 femoral head with that of a modern human necessitates all six DOFs to be included in which AL 288-1 could osteologically extend the hip to facilitate a human-like gait., This research was supported by a Leverhulme Trust Early Career Fellowship (Grant Number: ECF-2021-054) awarded to ALAW.

In this study, we want to investigate the covariation in the shape of two carpal bones, the triquetrum and hamate, and the possible association with locomotor behavior in a broad range of primate taxa. We applied 3D Geometric Morphometrics on a large data set comprising 309 anthropoid primates of 12 different genera. Principal component analyses were performed on the covariance matrix of 18 (triquetrum) and 23 (hamate) Procrustes-aligned surface landmarks. A two-block partial least square analysis was done to test the covariance between triquetrum and hamate shape, without relying on the predictive models implicit in regression analyses. The results show that the carpal shape of quadrupedal anthropoids, which mainly use their wrist under compressive conditions, differs from that of suspensory primates as their wrist is possibly subjected to tensile and torsional forces. Within the hominids, differences in shape also distinguish more terrestrial from more arboreal species. Even within the great apes, we are able to capture shape differences between species of the same genus. In combination with behavioral and biomechanical studies, the results of this research can be used to establish form-function relationships of the primate hand which will aid the functional interpretation of primate fossil remains.

Objectives Age-degenerative features of the metatarsals are poorly known despite the importance of metatarsal bone properties for investigating mobility patterns. We assessed the role of habitual activity in shaping the patterning and magnitude of sexual dimorphism in age-related bone loss in the hallucal metatarsal. Materials and methods Cross-sections were extracted at midshaft from micro-computed tomography scan models of individuals from medieval rural (Abingdon Vineyard) and early industrial urban (Spitalfields) settings (n = 71). A suite of cross-sectional geometry dimensions and biomechanical properties were compared between populations. Results The rural group display generally stronger and larger metatarsals that show a greater capacity to resist torsion and that have comparatively greater bending strength along the medio-lateral plane. Men in both groups show greater values of cortical area than women, but only in the urban group do men show lower magnitudes of age-related decline compared to females. Women in rural and urban populations show different patterns of age-related decline in bone mass, particularly old women in the urban group show a marked decline in cortical area that is absent for women in the rural group. Discussion Lifetime exposure to hard, physical activity in an agricultural setting has contributed to the attainment of greater bone mass and stronger bones in young adults. Furthermore, over the life-course, less of this greater amount of bone is lost, such that sustained activity levels may have acted to buffer against age-related decline, and this is most pronounced for women, who are expected to experience greater bone loss later in life than men.

Formby Point, England is a well-documented exposure site in which marine erosion has regularly exposed Neolithic human trackways along the coastline since the 1980s. We report here the discovery of an additional 17 trackways and 61 isolated tracks (181 human footprints in total) discovered during four field seasons of natural site exposure at four localities in the Formby Point region, recorded in 2016-2018. This adds substantially to the existing ichnological and palaeoecological body of data available for this site. The footprints belong to adults and children, in association with a large collection of other animal prints (>700), and are typically preserved in sandy-silts. The human trackways show a bimodal direction, with most of the trackmakers travelling in a south-west direction, towards the palaeo-coastline, with fewer trackmakers travelling in a north east direction, inland. Some trackmakers were either walking side by side or were following one another hours/days later as indicated by parallel trackways, whilst one trackway shows an individual running, eventually coming to a stop with both feet together on the ground. The trackmakers made repeated visits to the site, whereby some trackways were made hours/days earlier than others, evidenced by the impressions’ depth. We present the data here to: (1) add to literature of prehistoric footprints discovered around the world; and (2) make all three dimensional models and associated metadata publicly and freely available for use by other researchers. Most importantly, Formby is an exposure site in which we exclusively rely upon erosion to expose the footprints. Therefore, researchers can only understand the true dynamics of the site via the reporting of each successive exposure. We encourage local researchers to work with the public to aid in future documentation of the site to add to the continuously-growing database of Formby discoveries.

Elucidating when Neanderthal populations disappeared from Eurasia is a key question in paleoanthropology, and Belgium is one of the key regions for studying the Middle to Upper Paleolithic transition. Previous radiocarbon dating placed the Spy Neanderthals among the latest surviving Neanderthals in Northwest Europe with reported dates as young as 23,880 ± 240 B.P. (OxA-8912). Questions were raised, however, regarding the reliability of these dates. Soil contamination and carbon-based conservation products are known to cause problems during the radiocarbon dating of bulk collagen samples. Employing a compound-specific approach that is today the most efficient in removing contamination and ancient genomic analysis, we demonstrate here that previous dates produced on Neanderthal specimens from Spy were inaccurately young by up to 10,000 y due to the presence of unremoved contamination. Our compound-specific radiocarbon dates on the Neanderthals from Spy and those from Engis and Fonds-de-Forêt demonstrate that they disappeared from Northwest Europe at 44,200 to 40,600 cal B.P. (at 95.4% probability), much earlier than previously suggested. Our data contribute significantly to refining models for Neanderthal disappearance in Europe and, more broadly, show that chronometric models regarding the appearance or disappearance of animal or hominin groups should be based only on radiocarbon dates obtained using robust pretreatment methods.

Elucidating when Neanderthal populations disappeared from Eurasia is a key question in paleoanthropology, and Belgium is one of the key regions for studying the Middle to Upper Paleolithic transition. Previous radiocarbon dating placed the Spy Neanderthals among the latest surviving Neanderthals in Northwest Europe with reported dates as young as 23,880 ± 240 B.P. (OxA-8912). Questions were raised, however, regarding the reliability of these dates. Soil contamination and carbon-based conservation products are known to cause problems during the radiocarbon dating of bulk collagen samples. Employing a compound-specific approach that is today the most efficient in removing contamination and ancient genomic analysis, we demonstrate here that previous dates produced on Neanderthal specimens from Spy were inaccurately young by up to 10,000 y due to the presence of unremoved contamination. Our compound-specific radiocarbon dates on the Neanderthals from Spy and those from Engis and Fonds-de-Foret demonstrate that they disappeared from Northwest Europe at 44,200 to 40,600 cal B.P. (at 95.4% probability), much earlier than previously suggested. Our data contribute significantly to refining models for Neanderthal disappearance in Europe and, more broadly, show that chronometric models regarding the appearance or disappearance of animal or hominin groups should be based only on radiocarbon dates obtained using robust pretreatment methods.

Fossil hominin footprints provide a direct source of evidence of locomotor behavior and allow inference of other biological data such as anthropometrics. Many recent comparative analyses of hominin footprints have used 3D analytical methods to assess their morphological affinities, comparing tracks from different locations and/or time periods. However, environmental conditions can sometimes preclude 3D digital capture, as was the case at Happisburgh (England) in 2013. Consequently, we use here a 2D geometric morphometric approach to investigate the evolutionary context of the Happisburgh tracks. The comparative sample of hominin tracks comes from eight localities that span a broad temporal range from the Pliocene to Late Holocene. The results show disparity in the shapes of tracks ascribed to hominins from the Pliocene (presumably Australopithecus afarensis), Pleistocene (presumably Homo erectus and Homo antecessor), and Holocene (Homo sapiens). Three distinct morphological differences are apparent between time samples: changes in adduction of the hallux, changes in the shape and position of the medial longitudinal arch impression, and apparent changes in foot proportions. Linear dimensions classified the potential H. antecessor tracks from Happisburgh as being most similar to the presumed H. erectus prints from Ileret. We demonstrate using 2D geometric morphometric methods and linear dimensions that the Happisburgh tracks are morphologically similar to other presumed Homo tracks and differ from the Laetoli footprints. The probable functional implications of these results fit well with previous comparative analyses of hominin tracks at other sites.

The position (FMP) and orientation (FMO) of the foramen magnum have been used as proxies for locomotion and posture in extant and extinct primates. Several indices have been designed to quantify FMP and FMO but their application has led to conflicting results. Here, we test six widely used indices and two approaches (univariate and multivariate) for their capability to discriminate between postural and locomotor types in extant primates and fossil hominins. We then look at the locomotion of australopithecines and Homo on the base of these new findings. The following measurements are used: the opisthocranion-prosthion (OP-PR) and the opisthocranion-glabella (OP-GL) indices, the basion-biporion (BA-BP) and basion-bicarotid chords, the foramen magnum angle (FMA), and the basion-sphenoccipital ratio. After exploring the indices variability using principal component analysis, pairwise comparisons are performed to test for the association between each index and the locomotor and postural habits. Cranial size and phylogeny are taken into account. Our analysis indicates that none of the indices or approaches provides complete discrimination across locomotor and postural categories, although some differences are highlighted. FMA and BA-BP distinguish respectively obligate and facultative bipeds from all other groups. For what concerns posture, orthogrades and pronogrades differ with respects to OP-PR, OP-GL, and FMA. Although the multivariate approach seems to have some discrimination power, the results are most likely driven by facial and neurocranial variability embedded in some of the indices. These results demonstrate that indices relying on the anteroposterior positioning of the foramen may not be appropriate proxies for locomotion among primates. The assumptions about locomotor and postural habits in fossil hominins based on foramen magnum indices should be revised in light of these new findings.

There is evidence for early Pleistocene Homo in northern Europe, a novel hominin habitat. Adaptations enabling this colonisation are intriguing given suggestions that Homo exhibits physiological and behavioural malleability associated with a ‘colonising niche’. Differences in body size/shape between conspecifics from different climates are well-known in mammals, could relatively flexible size/shape have been important to Homo adapting to cold habitats? If so, at what point did this evolutionary stragegy arise? To address these questions a base-line for adaptation to climate must be established by comparison with outgroups. We compare skeletons of Japanese macaques from four latitudes and find inter-group differences in postcranial and cranial size and shape. Very small body mass and cranial size in the Southern-most (island) population are most likely affected by insularity as well as ecogeographic scaling. Limb lengths and body breadths show group differences that accord with the expectations of thermoregulation across the whole range of latitudes. Postcranial size appears to vary more than shape, yet there is also evidence that limb segments follow Allen's rule in the forelimb at least, suggesting differing climatic signals in different regions of the skeleton. In contrast to other intraspecific studies of catarrhine ecogeography, the results presented here demonstrate non-allometric latitudinal patterns in craniofacial shape in Japanese macaques, which align closely with what is seen in cold-adapted humans. These insights begin to provide a comparison for hominin adaptation to similar habitat diversity and the role of biological adaptation in shaping the evolution and dispersal of Homo species.

Introduction Fossilised footprints have been discovered all over the world and can provide information regarding the foot size and subsequent body size estimates of the track makers or an insight into the kinematics of the foot/lower limb. After exposure, these fossils rapidly erode. It is predicted that footprint morphology is compromised after creation, prior to fossilisation and that erosion after exposure will affect the morphology of a footprint after exposure. To date, no studies have assessed if degradation prior to fossilisation and/or after fossilisation, and subsequent exposure, affects the morphology of the print, thereby affecting any measurements taken. This study aims to quantify these pre- and post-erosional processes. Materials and methods A set of experimentally generated footprints were created to test the effects of degradation of footprint morphology prior to fossilisation. In addition, Holocene footprints were recorded at Formby Point, Sefton, UK. In just over a week tidal action had completely eroded the Holocene beds. Photogrammetry was applied to the experimental human footprints and a selection of Holocene human and animal footprints. Three-dimensional Geometric Morphometric methods were utilised to estimate differences in shape and size. Results Results from the experimental footprints indicate that weather action affects the size and shape of a footprint prior to fossilisation. When the weather was dry, footprint shape and size showed little difference for two weeks, but rainfall caused significant changes. The Holocene footprints show that after fossilisation and exposure to coastal erosion, footprint rigidity is highly compromised. The human footprint borders progressively recede, increasing length and width each day. Footprint depth, often used to inform upon speed and kinematics, varied considerably in one week. Some regions becoming shallower, others increasing in depth. Similar results were found for the animal footprints, but with less significant changes in shape and size determined. Conclusion Observed significant differences in measurements result in problems for predicting stature, mass, sex, and kinematic analyses. This warrants caution when making interpretations from fossilised footprints. Rapid recording of fossilised prints from first exposure and assessing pre-fossilisation processed are necessities when recording footprint surfaces.

Objectives: A variety of mechanical processes can result in ante-mortem dental chipping. In this study, chipping data in the teeth of Homo naledi are compared with those of other pertinent dental samples to give insight into their etiology. Materials and Methods: Permanent teeth with complete crowns evidencing occlusal wear were examined macroscopically. The location, number, and severity of fractures were recorded and compared to those found in samples of two other South African fossil hominin species, as well as in samples of non-human primates (n= 3) and recent humans (n= 7). Results: With 44% of teeth affected, Homo naledi exhibits far higher rates of chipping than the other fossil hominin samples. Specifically, 50% of posterior teeth and 31% of anterior teeth display at least one chip. The maxillary teeth are more affected than the mandibular teeth (45% vs. 43%, respectively), 73% of molar chipping occurs on interproximal surfaces, and right teeth are more often affected than left teeth (50% vs. 38%). Discussion: Results indicate that the teeth of H. naledi were exposed to acute trauma on a regular basis. Because interproximal areas are more affected than buccal, and posterior teeth more than anterior, it is unlikely that non-masticatory cultural behavior was the cause. A diet containing hard and resistant food, or contaminants such as grit, is more likely. The small chip size, as well as steep occlusal wear and cupped dentine on some molars is supportive of the latter possibility. This pattern of chipping suggests H. naledi differed considerably – in terms of diet, environment, and/or specialized masticatory processing— relative to other African fossil hominins.

Caries frequencies in South African fossil hominins were observed and compared with other hominin samples. Species studied includeParanthropus robustus,Homo naledi,Australopithecus africanus,early HomoandA. sediba. Teeth were viewed macroscopically with Micro-CT scans used to confirm lesions. Position and severity of each lesion were also noted and described. For all South African fossil hominin specimens studied, 16 have carious lesions, six of which are described for the first time in this study. These are from a minimum of six individuals, and include fourP. robustus,oneH. naledi,and oneearly Homoindividual. No carious lesions were found on deciduous teeth, or any teeth assigned toA. africanus. Most are located interproximal, and only posterior teeth are affected. Caries frequency typically ranges between 1-5% of teeth in non-agricultural human samples, and this pattern seemingly holds true for at least the past two million years in the hominin lineage. Non-agricultural populations significantly above or below this threshold generally have a specialized diet, supporting other dietary evidence thatA. africanuslikely consumed large amounts of tough, non-cariogenic vegetation. Given the common occurrence of caries in the other hominin species, cariogenic bacteria and foods were evidently common in their collective oral environment. Along with recent research highlighting additional examples of caries inH. neanderthalensis, earlyHomoand PleistoceneH. sapiens, caries is clearly an ancient disease that was much more common than once maintained throughout the course of human evolution.

The construction of the biological profile from human skeletal remains is the foundation of anthropological examination. However, remains may be fragmentary and the elements usually employed, such as the pelvis and skull, are not available. The clavicle has been successfully used for sex estimation in samples from Iran and Greece. In the present study, the aim was to test the suitability of the measurements used in those previous studies on a British Medieval population. In addition, the project tested whether discrimination between sexes was due to size or clavicular strength. The sample consisted of 23 females and 25 males of pre-determined sex from two medieval collections: Poulton and Gloucester. Six measurements were taken using an osteometric board, sliding calipers and graduated tape. In addition, putty rings and bi-planar radiographs were made and robusticity measures calculated. The resulting variables were used in stepwise discriminant analyses. The linear measurements allowed correct sex classification in 89.6% of all individuals. This demonstrates the applicability of the clavicle for sex estimation in British populations. The most powerful discriminant factor was maximum clavicular length and the best combination of factors was maximum clavicular length and circumference. This result is similar to that obtained by other studies. To further investigate the extent of sexual dimorphism of the clavicle, the biomechanical properties of the polar second moment of area J and the ratio of maximum to minimum bending rigidity are included in the analysis. These were found to have little influence when entered into the discriminant function analysis.

This paper assesses the earliest evidence for widespread dental modification in Northwest Africa. The intentional modification of teeth has implications for an individual's appearance, sense of identity and perceived status. The range of modifications reported varies from alterations of shape or color to the complete removal of healthy teeth (evulsion or ablation). The availability of well-dated collections reveals that Northwest Africa was the first region where the custom of tooth evulsion was widely practiced. Analysis of Iberomaurusian (Late Stone Age, n = 77) and Capsian dental material (n = 12) shows that evulsion was present in most male and female individuals (>94%). The most common Iberomaurusian practice involved removal of both upper central incisors (around 65%) although removal of fewer, none and more teeth was also recorded. Observations of the extent of alveolar remodeling of different sockets revealed that teeth were frequently removed at different ages, suggesting that the cultural significance was age transgressive and may have related to an event that individuals experienced more than once. During the Capsian period the prevalence of evulsion was lower in males than in females, but when present more teeth were removed with evulsion frequently involving both mandible and maxilla. Tooth wear analysis shows that evulsion affected not only the appearance of the individual but also the functioning of the masticatory complex.

Hominin fossilised trackways are commonly used to reconstruct locomotory behaviour and to characterise track-maker biometrics. They are the most direct representation of hominin locomotion available, yet the fossil sites are susceptible to extreme cases of erosion and weathering, often resulting in the destruction of the fossil beds, as was the case at two prehistoric footprint sites: Formby Point and Happisburgh, UK. Footprints can also be found in easily deformable sediments where the excavator can unintentionally trample and destroy the prints, as occurred at Formby Point, UK. There is the need for non-destructive methods to be identified that can rapidly record the fossil area before further damage can occur to the fossil beds. Because current methods of 3D data capture (laser scanning or photogrammetry) can be invasive (i.e., the excavator is often required to trample the sediment layer during data capture), non-invasive methods need to be explored to circumvent issues in advertently destroying fossil material. In this project, a series of experiments using non-invasive methods tested the applicability of Unmanned Aerial Vehicle (UAV) technology combined with photogrammetry to rapidly and accurately record footprints before further damage to the fossil interface occurred. Various flight paths, UAVs, cameras and capture types (video versus camera stills) were incorporated to test the accuracy in minute depth reconstruction and subsequent 3D mesh creation of small, intricate detail. Shape and size, the consequence of poorly reconstructed depth dimensionality of surfaces, were found to be affected by flight path and by the height of the UAV, indicating that the most optimal method of recording tracks is to use a handheld DSLR camera following a circular or rastered flight path. Despite attempts to utilise UAV technology, inaccurate reconstructions of footprint topography were produced. Although UAV technology produced unreliable reconstructions, UAVs remain a technological solution when sites may be at immediate risk of destruction, as was the case at Happisburgh, UK. The deployment of UAV technology will permit the digital preservation of fossil material which would otherwise be lost.

The reduction in dental size and mandibular robusticity is regarded as a major trend in human evolution, traditionally considered the result of the peculiar extra-oral food processing skills of Homo. The use of stone tools and fire would have allowed our ancestors to chew softer food in smaller bite size, thus relaxing the selective pressures to keep a large dentition and a robust lower jaw. This perspective assumes that differences in dental size and mandibular robusticity in hominins represent functional dissimilarities. This study uses a catarrhine comparative approach to test this fundamental assumption of the hypotheses on dental and mandibular reduction in Homo. A sample of extant catarrhines and fossil hominins was used to test for correlations between dental size, mandibular robusticity, and dietary proxies, the latter include diet quality, diet heterogeneity, feeding time, and microwear variables. The effects of phylogeny and body size were considered. Findings support the association between technological developments in Homo and reduction in incisor size and mandibular corpus robusticity, though not for premolar, molar size, and symphyseal robusticity. These results challenge the functional interpretation of postcanine reduction and symphyseal changes in the genus Homo.

The taxonomic attribution of the skeletal material of Sima de los Huesos (SH, Spain) has puzzled researchersfor decades. At the time of their discovery, the SH hominins were considered as pre-Neanderthal based on their morphology [1]. Later, tooth dimensions were found to link the SH sample to middle Pleistocene specimens such as Mauer, Montmaurin, and Arago II [2], which are commonly ascribed to Homo heidelbergensis. More recently, a close relationship between Neanderthals and the SH specimens was suggested based on the morphology of cranial vault and face, in particular regarding the masticatory apparatus [3]. Decades of unresolved debate produced three plausible scenarios for the status of SH in the human ancestry: (I) SH has to be regarded as part of the variability of H. heidelbergensis, (II) SH should be considered H. neanderthalensis, (III) SH andH. heidelbergensiscould represent two separate evolutionary lineages co-existing during middle Pleistocene in Eurasia, with SH being phylogenetically closer to Neanderthals. In this work, we applied supervised machine learning to test for the affinity of SH (N: 126) to H. heidelbergensis (N: 13), H. neanderthalensis (N: 73), and H. sapiens (N: 403). The sample included Mesio-Distal and Bucco-Lingual diameters of mandibular postcanine dentition. Because of the fragmentary nature of the dental fossil and archaeological records, several missing data were included in the dataset and were estimated using Multiple Imputation via Predictive Mean Matching [4]. To avoid biases due to disproportions in sample size, missing data were estimated separately for each group considered. A recursive partitioning algorithm known as Conditional Inference Tree (CIT) [5] was used to build a hierarchical model for predicting the classification of dental metric data into each of the categorical groups considered (SH, H. heidelbergensis, H. neanderthalensis, and H. sapiens). The data was divided in one training set and one test set. The training set (N: 105) was used to allow the CIT algorithm to determine the best classification solution for the dental data. The test set (N: 497) was necessary to control the accuracy of classification. Our results show that when SH is considered as part of H. heidelbergensis it is easily misclassified as Neanderthal (33.6% accuracy). On the other side, SH is recognised as an independent group when other H. heidelbergensis specimens are discarded from the analysis (95.6% accuracy). In both cases, the classification is based on a major contribution of the Bucco-Lingual diameter of the third premolar in association with the dimensions of the second and third molars. The accuracy associated with the H. sapiens and H. neanderthalensis samples indicates that the model is capable of classifying discrete taxonomic groups. Our results support the hypothesis that SH represents a distinct evolutionary lineage from H. heidelbergensis and suggest that SH could be closely related to “classic” Neanderthals. This work represents the first application of supervised machine learning to Palaeoanthropology and highlights the potential of this field to the study of human evolution. References:[1] Aguirre, E., & De Lumley, M. A. (1977). Fossil men from Atapuerca, Spain: their bearing on human evolution in the Middle Pleistocene. Journal of human evolution, 6(8), 681-688.[2] de Castro, J. M. B. (1986). Dental remains from Atapuerca (Spain) I. Metrics. Journal of Human Evolution, 15(4), 265-287.[3] Arsuaga, J. L., Martínez, I., Arnold, L. J., Aranburu, A., Gracia-Téllez, A., Sharp, W. D., ... & Poza-Rey, E. (2014). Neandertal roots: Cranial and chronological evidence from Sima de los Huesos. Science, 344(6190), 1358-1363.[4] Morris, T. P., White, I. R., & Royston, P. (2014). Tuning multiple imputation by predictive mean matching and local residual draws. BMC medical research methodology, 14(1), 75.[5] Strobl, C., Malley, J., & Tutz, G. (2009). An introduction to recursive partitioning: rationale, application, and characteristics of classification and regression trees, bagging, and random forests. Psychological methods, 14(4), 323

In 1884, Arthur Smith Woodward first met Charles Dawson, a solicitor and industrious amateur collector, antiquarian, geologist, archaeologist and palaeontologist. This began a long association and friendship centred on their mutual interest in palaeontology and human evolution. Dawson devised a complicated plot focused around the ancient river gravel deposits at Barkham Manor near the village of Piltdown, Sussex. In these gravels he planted stone tools and fossil mammal remains together with the lower jaw of an ape and numerous modern human cranial bones to deceive the scientific establishment into believing an early human ancestor had been found in his own back yard. Cleverly devised to provide anatomists and archaeologists with evidence for concepts that they wanted to believe were true, Dawson fuelled numerous contentious debates among scientists that quickly attracted international attention. Nothing could be more unfortunate than such a respectable scientist as Arthur Smith Woodward being taken in by the events of 1912, and then subsequently swept along by them well into his retirement right up to the time of his death in 1944.

This chapter described the earliest evidence of the systematic practice of ablation. Purposeful removal of the upper central incisors became a widespread practice with the Iberomaurusian of the Later Stone Age in the Maghreb region—the area of current-day northern Morocco, Algeria, and Tunisia. During the Capsian times, ablation became less prevalent, but in some cases all incisors and canines were removed. During the Neolithic period the practice became even less frequent and in some areas disappeared completely. Nevertheless, in some regions of North Africa ablation remained common and may have been a way of identifying certain tribes or individuals within society. This chapter also considers possible causes for the origin of the practice.

In 2001, a collection of skeletal material was donated to the Natural History Museum, London, by the Royal College of Surgeons, London. It consisted of boxes discovered among the personal belongings of Sir Arthur Keith. This paper describes the work undertaken to identify and document the human skeletal material in the Keith Collection. The study identified the human fossils as having come from a number of excavations directed by Dorothy Garrod in the 1920s and 1930s in Israel. The collection contains the long considered lost human skeletal collection from the type-site of the Natufian industry: Shukbah Cave. The majority of this material is of Natufian origin but contains a few Neanderthal specimens. A small amount of heavily fragmented bones associated with Skhūl VII and IX were also found. The most remarkable of the re-discovered collection is the material from el-Wad and Kebara Caves. It was identified to be the missing material from the Middle and Upper Paleolithic levels briefly described in 1939 in The Stone Age of Mount Carmel vol 2. by Theodore McCown and Sir Arthur Keith. These important fossils hold great potential to answer questions about the Middle to Upper Paleolithic transition in the Near East, and the emergence of anatomically modern humans.

Dental caries is an infectious disease that causes tooth decay. The high prevalence of dental caries in recent humans is attributed to more frequent consumption of plant foods rich in fermentable carbohydrates in food-producing societies. The transition from hunting and gathering to food production is associated with a change in the composition of the oral microbiota and broadly coincides with the estimated timing of a demographic expansion in Streptococcus mutans, a causative agent of human dental caries. Here we present evidence linking a high prevalence of caries to reliance on highly cariogenic wild plant foods in Pleistocene hunter-gatherers from North Africa, predating other high caries populations and the first signs of food production by several thousand years. Archaeological deposits at Grotte des Pigeons in Morocco document extensive evidence for human occupation during the Middle Stone Age and Later Stone Age (Iberomaurusian), and incorporate numerous human burials representing the earliest known cemetery in the Maghreb. Macrobotanical remains from occupational deposits dated between 15,000 and 13,700 cal B.P. provide evidence for systematic harvesting and processing of edible wild plants, including acorns and pine nuts. Analysis of oral pathology reveals an exceptionally high prevalence of caries (51.2% of teeth in adult dentitions), comparable to modern industrialized populations with a diet high in refined sugars and processed cereals. We infer that increased reliance on wild plants rich in fermentable carbohydrates and changes in food processing caused an early shift toward a disease-associated oral microbiota in this population.

A variety of mechanical processes can result in antemortem dental chipping. In this study, chipping data in the teeth of Homo naledi are compared with those of other pertinent dental samples to give insight into their etiology.Permanent teeth with complete crowns evidencing occlusal wear were examined macroscopically. The location, number, and severity of fractures were recorded and compared to those found in samples of two other South African fossil hominin species and in samples of nonhuman primates (n = 3) and recent humans (n = 7).With 44% of teeth affected, H. naledi exhibits far higher rates of chipping than the other fossil hominin samples. Specifically, 50% of posterior teeth and 31% of anterior teeth display at least one chip. The maxillary teeth are more affected than the mandibular teeth (45% vs 43%, respectively), 73% of molar chipping occurs on interproximal surfaces, and right teeth are more often affected than left teeth (50% vs 38%).Results indicate that the teeth of H. naledi were exposed to acute trauma on a regular basis. Because interproximal areas are more affected than buccal and posterior teeth more than anterior, it is unlikely that nonmasticatory cultural behavior was the cause. A diet containing hard and resistant food, or contaminants such as grit, is more likely. The small chip size, and steep occlusal wear and cupped dentine on some molars are supportive of the latter possibility. This pattern of chipping suggests that H. naledi differed considerably-in terms of diet, environment, and/or specialized masticatory processing-relative to other African fossil hominins.

Organic (bone, antler, wood) knapping tools were undoubtedly a component of early human tool kits since the Lower Palaeolithic. Previous studies have identified pitting and the occasional presence of embedded flint flakes as important features for recognizing archaeological bone and antler percussors. However, no systematic protocol of analysis has been suggested for the study of this rare archaeological material. Here we present qualitative and quantitative results of a preliminary analysis of an experimental knapping hammer, using a novel combination of microscopy (focus variation optical microscope and scanning electron microscope), micro-CT scanning and energy dispersive X-ray spectroscopy. These imaging and analytical techniques are used to characterize use-damage from the manufacture of handaxes. This paper highlights the strengths and weakness of each technique. Use-wear on the working area included attritional bone loss, micro-striations and compaction of the outer layer of the antler matrix from repeated hitting of the beam against the sharp edge of the handaxe during knapping. Embedded flint flakes were also identified in the pits and grooves. This combination of high-resolution imaging techniques is applicable to fragile archaeological specimens, including those encrusted by sediment or encased in matrix.

In 1912, palaeontologist Arthur Smith Woodward and amateur antiquarian and solicitor Charles Dawson announced the discovery of a fossil that supposedly provided a link between apes and humans: Eoanthropus dawsoni (Dawson's dawn man). The publication generated huge interest from scientists and the general public. However, ‘Piltdown man's’ initial celebrity has long been overshadowed by its subsequent infamy as one of the most famous scientific frauds in history. Our re-evaluation of the Piltdown fossils using the latest scientific methods (DNA analyses, high-precision measurements, spectroscopy and virtual anthropology) shows that it is highly likely that a single orang-utan specimen and at least two human specimens were used to create the fake fossils. The modus operandi was found consistent throughout the assemblage (specimens are stained brown, loaded with gravel fragments and restored using filling materials), linking all specimens from the Piltdown I and Piltdown II sites to a single forger—Charles Dawson. Whether Dawson acted alone is uncertain, but his hunger for acclaim may have driven him to risk his reputation and misdirect the course of anthropology for decades. The Piltdown hoax stands as a cautionary tale to scientists not to be led by preconceived ideas, but to use scientific integrity and rigour in the face of novel discoveries.

The research was funded by the Calleva Foundation as part of the Pathways to Ancient Britain Project. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Investigations at Happisburgh, UK, have revealed the oldest known hominin footprint surface outside Africa at between ca. 1 million and 0.78 million years ago. The site has long been recognised for the preservation of sediments containing Early Pleistocene fauna and flora, but since 2005 has also yielded humanly made flint artefacts, extending the record of human occupation of northern Europe by at least 350,000 years. The sediments consist of sands, gravels and laminated silts laid down by a large river within the upper reaches of its estuary. In May 2013 extensive areas of the laminated sediments were exposed on the foreshore. On the surface of one of the laminated silt horizons a series of hollows was revealed in an area of ca. 12 m2. The surface was recorded using multi-image photogrammetry which showed that the hollows are distinctly elongated and the majority fall within the range of juvenile to adult hominin foot sizes. In many cases the arch and front/back of the foot can be identified and in one case the impression of toes can be seen. Using foot length to stature ratios, the hominins are estimated to have been between ca. 0.93 and 1.73 m in height, suggestive of a group of mixed ages. The orientation of the prints indicates movement in a southerly direction on mud-flats along the river edge. Early Pleistocene human fossils are extremely rare in Europe, with no evidence from the UK. The only known species in western Europe of a similar age is Homo antecessor, whose fossil remains have been found at Atapuerca, Spain. The foot sizes and estimated stature of the hominins from Happisburgh fall within the range derived from the fossil evidence of Homo antecessor. Publisher PDF

One of the author names was misspelled in the paper. ‘Lucia Burgia’ should be amended to ‘Lucia Burgio’ and the author list should be as follows

Archaeological assemblages often lack the complete long bones needed to estimate stature and body mass. The most accurate estimates of body mass and stature are produced using femoral head diameter and femur length. Foot bones including the first metatarsal preserve relatively well in a range of archaeological contexts. In this article we present regression equations using the first metatarsal to estimate femoral head diameter, femoral length, and body mass in a diverse human sample. The skeletal sample comprised 87 individuals (Andamanese, Australasians, Africans, Native Americans, and British). Results show that all first metatarsal measurements correlate moderately to highly (r = 0.62-0.91) with femoral head diameter and length. The proximal articular dorsoplantar diameter is the best single measurement to predict both femoral dimensions. Percent standard errors of the estimate are below 5%. Equations using two metatarsal measurements show a small increase in accuracy. Direct estimations of body mass (calculated from measured femoral head diameter using previously published equations) have an error of just over 7%. No direct stature estimation equations were derived due to the varied linear body proportions represented in the sample. The equations were tested on a sample of 35 individuals from Christ Church Spitalfields. Percentage differences in estimated and measured femoral head diameter and length were less than 1%. This study demonstrates that it is feasible to use the first metatarsal in the estimation of body mass and stature. The equations presented here are particularly useful for assemblages where the long bones are either missing or fragmented, and enable estimation of these fundamental population parameters in poorly preserved assemblages.

Since their discovery, Neanderthals have been described as having a marked degree of anteroposterior curvature of the femoral shaft. Although initially believed to be pathological, subsequent discoveries of Neanderthal remains lead femoral curvature to be considered as a derived Neanderthal feature. A recent study on Neanderthals and middle and early Upper Palaeolithic modern humans found no differences in femoral curvature, but did not consider size-corrected curvature. Therefore, the objectives of this study were to use 3D morphometric landmark and semi-landmark analysis to quantify relative femoral curvature in Neanderthals, Upper Palaeolithic and recent modern humans, and to compare adult bone curvature as part of the overall femoral morphology among these populations. Comparisons among populations were made using geometric morphometrics (3D landmarks) and standard multivariate methods. Comparative material involved all available complete femora from Neanderthal and Upper Palaeolithic modern human, archaeological (Mesolithic, Neolithic, Medieval) and recent human populations representing a wide geographical and lifestyle range. There are significant differences in the anatomy of the femur between Neanderthals and modern humans. Neanderthals have more curved femora than modern humans. Early modern humans are most similar to recent modern humans in their anatomy. Femoral curvature is a good indicator of activity level and habitual loading of the lower limb, indicating higher activity levels in Neanderthals than modern humans. These differences contradict robusticity studies and the archaeological record, and would suggest that femoral morphology, and curvature in particular, in Neanderthals may not be explained by adult behavior alone and could be the result of genetic drift, natural selection or differences in behavior during ontogeny.

Here we describe and evaluate a new method for quantifying long bone curvature using geometric morphometric and semi-landmark analysis of the human femur. The technique is compared with traditional ways of measuring subtense and point of maximum curvature using either coordinate calipers or projection onto graph paper. Of the traditional methods the graph paper method is more reliable than using coordinate calipers. Measurement error is consistently lower for measuring point of maximum curvature than for measuring subtense. The results warrant caution when comparing data collected by the different traditional methods. Landmark data collection proves reliable and has a low measurement error. However, measurement error increases with the number of semi-landmarks included in the analysis of curvature. Measurements of subtense can be estimated more reliably using 3D landmarks along the curve than using traditional techniques. We use equidistant semi-landmarks to quantify the curve because sliding the semi-landmarks masks the curvature signal. Principal components analysis of these equidistant semi-landmarks provides the added benefit of describing the shape of the curve. These results are promising for functional and forensic analysis of long bone curvature in modern human populations and in the fossil record.

Objectives: Although the evolution of the hominin masticatory apparatus has been linked to diet and food processing, the physical connection between neurocranium and lower jaw suggests a role of encephalization in the trend of dental and mandibular reduction. Here, the hypothesis that tooth size and mandibular robusticity are influenced by morphological changes in the neurocranium was tested. Materials and Methods: Three-dimensional landmarks, alveolar lengths, and mandibular robusticity data were recorded on a sample of chimpanzee and human skulls. The morphological integration between the neurocranium and the lower jaw was analyzed by means of Singular Warps Analysis. Redundancy Analysis was performed to understand if the pattern of neuromandibular integration affects tooth size and mandibular robusticity. Results: There is significant morphological covariation between neurocranium and lower jaw in both chimpanzees and humans. In humans, changes in the temporal fossa seem to produce alterations of the relative orientation of jaw parts, while the influence of similar neurocranial changes in chimpanzees are more localized. In both species, postcanine alveolar lengths and mandibular robusticity are associated with shape changes of the temporal fossa. Conclusions: The results of this study support the hypothesis that the neurocranium is able to affect the evolution and development of the lower jaw, although most likely through functional integration of mandible, teeth, and muscles within the masticatory apparatus. This study highlights the relative influence of structural constraints and adaptive factors in the evolution of the human skull.

© 2018 Elsevier Ltd The supposed herbivorous cave bear (Ursus spelaeus) occupied Europe throughout the Quaternary. Being subject to large spatial variation has led to the intensive study on its geographical polymorphism, generating debates on sub-speciation. However, temporal morphological information on the species is somewhat lacking. Here, we apply geometric morphometrics (GMM) technique to investigate temporal morphological variation in molar size and shape of Ursus spelaeus from different chronostratigraphic sediment units in a geographically confined site (Scladina Cave, Belgium), covering approximately 100,000 years. Our findings show significant morphological variation between groups analysed in both size and shape. M2 shows a chronological size increase with PCA plots visually expressing differences in all groups, relating to a buccolingual expansion and an increase of the talon masticatory platform through time. Reduction in the M1 is also shown, possibly to maintain biomechanical performance of dentition for effective mastication, more so in groups relating to the latter stages of the Quaternary. Findings suggest a rapid response to climatic factors constraining consumable food sources, with GMM offering a promising analytical approach in understanding the palaeobiology, palaeoecology and morphological variation in extinct and extant fossil mammals.

Out of all extinct megafaunal mammals of the Quaternary, the cave bear Ursus spelaeus is one of the best represented in the fossil record. This species has been found to exhibit skeletal morphological adaptations when exploiting varied environmental niches, be that spatially or temporally. Here, we employ geometric morphometrics and phenotypic trajectory analysis to explore temporal morphological changes across the entire lower molar tooth row from the infilling of Scladina Cave, Belgium. We show that molar tooth size increases from Marine Isotope Stage (MIS) 5 – MIS 3, with cusp position varying temporally in relation to a larger talonid grinding platform in later time periods. Phenotypic trajectory analyses further show similar evolutionary shape trajectories in the first and second molars, but not in the third molar. Morphological changes related to a larger grinding platform are found in the second and third molars, with the divergent morphological change of M3 suggesting that this tooth is less constrained and appears much more responsive to environmental changes. The need to cope with harder fibrous plant material present in the palaeoenvironment potentially constrained morphological evolution of the cave bear until its disappearance throughout Europe.

The use of soft (bone, antler, tooth and wood) hammers and retouchers is a key innovation in early stone tool technology, first appearing in the archaeological record with Lower Palaeolithic handaxe industries (e.g. Boxgrove, UK ~ 500 kya). Although organic knapping tools were undoubtedly a component of early human toolkits and are essential, for example, for the manufacture of finely-flaked handaxes, Mousterian scrapers and Upper Palaeolithic blades tools, such archaeological finds are exceptionally rare. In this study, we present qualitative and quantitative analyses (focus variation optical microscope, scanning electron microscope, micro-CT scanning and energy dispersive X-ray spectroscopy), to characterise use-damage on an antler base from Laugerie-Haute West (France). This specimen was originally identified as a waste-product from splinter manufacture, and the use-damage appears to have been missed. The new analysis shows that prior to being used as a flint-knapping percussor, the red deer antler had been further modified to reduce the length of its beam and to remove the tines. Although minimally used, characteristic use-damage includes attrition (pits and scores), compression of the antler matrix and flint chips embedded within some of the percussion features on the base of the burr. An AMS radiocarbon date of 12385 ± 55 BP (12647 ± 335 BC calibrated) confirms a Magdelanian context for the hammer. The fact that the Laugerie Haute knapping hammer went unrecognised in a well-studied and accessible collection for almost 200 years since its discovery, suggests that antler hammers may be more common than generally assumed. Only further re-examination of prehistoric antlers in museum collections will confirm whether the apparent rarity of antler hammers during the Lower and Middle Palaeolithic is real phenomenon or the result of analytical biases.

Funder: Liverpool John Moores University; Id: http://dx.doi.org/10.13039/501100004144, Estimation of stature from footprint lengths is a common prediction in forensic cases and in paleoanthropology upon the discovery of fossil footprints. Many studies, which have estimated stature from footprints, generally use a "one-size-fits-all" approach that usually involves applying a known ratio of foot length to total stature to do so, although this method has fallen out of practice in forensic cases in recent years but is still commonly used for fossil trace evidence. Yet, we know that substrate and speed can change the dimensions of a footprint, so why are these "one-size-fits-all" approaches still used today? We tested footprint production across different substrates at a walk, a fast walk, and a jog. We calculated how accurately footprint dimensions were impressed between these different conditions and identified sources of error in footprint lengths, and the percentage changes of how significantly a footprint can change in length between different conditions. We provide a table with different ratios that we encourage practitioners/field scientists to refer to and use when estimating stature from footprints, with respect to the substrate on which the footprint was created and the speed at which it was created. We actively encourage researchers to add the ratios by testing more substrates so that in the future stature can be more accurately estimated, thus aiding the paleoanthropological community, but also forensic investigations by statistically highlighting how different conditions can affect trace dimensions.