Ulnar shape of extant primates: Functional signals and covariation with triquetrum shape.

OBJECTIVES: In this study, we investigated the shape differences of the distal ulna in a phylogenetic context among a broad range of primate taxa. Furthermore, we evaluated covariation between ulnar and triquetrum shape and a possible association between ulnar shape and locomotor behavior. MATERIALS AND METHODS: We applied 3D geometric morphometrics on a large dataset comprising the distal ulna of 124 anthropoid primate specimens belonging to 12 different genera. For each species, a mean shape was calculated using 11 Procrustes-aligned surface landmarks on the distal ulna. These mean shapes are used in a bgPCA, pPCA, and PACA and 3D morphs were used to visualize more subtle differences between taxa. A p2B-PLS analysis was performed to test the covariance between distal ulnar and triquetrum shape. RESULTS: The results show that more closely related species exhibit a similar distal ulnar shape. Overall, extant hominid ulnae show a shape shift compared to those of extant monkeys and hylobatids. This includes a shortening of the ulnar styloid process and dorspalmarly widening of the ulnar head, shape characteristics that are independent of phylogeny. Within the hominids, Pongo pygmaeus seem to possess the most plesiomorphic distal ulnar shape, while Gorilla and Homo sapiens display the most derived distal ulna. Cercopithecoids, hylobatids, and P. pygmaeus are characterized by a relatively deep ECU groove, which is a shape trait dependent of phylogeny. Although there was no significant covariation between distal ulnar shape and triquetrum shape, the shape differences of the distal ulna between the different primate taxa reveal a possible link with locomotor behavior. CONCLUSIONS: The comparative analyses of this study reveal different shape trends in a phylogenetic context. Highly arboreal primates, such as hylobatids and Ateles fusciceps, show a distal ulnar morphology that appears to be adapted to tensile and torsional forces. In primates that use their wrist under more compressive conditions, such as quadrupedal cercopithecoids and great apes, the distal ulnar morphology seems to reflect increased compressive forces. In modern humans, the distal ulnar shape can be associated to enhanced manipulative skills and power grips. There was no significant covariation between distal ulnar shape and triquetrum shape, probably due to the variation in the amount of contact between the triquetrum and ulna. In combination with future research on wrist mobility in diverse primate taxa, the results of this study will allow us to establish form-function relationships of the primate wrist and contribute towards an evidence-based interpretation of fossil remains.

Principal component and linear discriminant analyses for the classification of hominoid primate specimens based on bone shape data.

In this study, we tested the hypothesis that machine learning methods can accurately classify extant primates based on triquetrum shape data. We then used this classification tool to observe the affinities between extant primates and fossil hominoids. We assessed the discrimination accuracy for an unsupervised and supervised learning pipeline, i.e. with principal component analysis (PCA) and linear discriminant analysis (LDA) feature extraction, when tasked with the classification of extant primates. The trained algorithm is used to classify a sample of known fossil hominoids. For the visualization, PCA and uniform manifold approximation and projection (UMAP) are used. The results show that the discriminant function correctly classified the extant specimens with an F1-score of 0.90 for both PCA and LDA. In addition, the classification of fossil hominoids reflects taxonomy and locomotor behaviour reported in literature. This classification based on shape data using PCA and LDA is a powerful tool that can discriminate between the triquetrum shape of extant primates with high accuracy and quantitatively compare fossil and extant morphology. It can be used to support taxonomic differentiation and aid the further interpretation of fossil remains. Further testing is necessary by including other bones and more species and specimens per species extinct primates.

Covariation between wrist bone morphology and maximal range of motion during ulnar deviation and supination in extant nonhuman primate taxa.

This study investigates the maximal range of motion (ROM) during wrist deviation and forearm rotation for five different primate genera and the possible correlation with the shape of the distal ulna, triquetrum and hamate. A two-block phylogenetic partial least square analysis was performed to test this covariation in a phylogenetic context, using shape coordinates and a matrix of maximal ROM data as input data. The results show that gibbons have the highest ROM for both ulnar deviation and supination, whereas Macaca exhibited the lowest ROM for supination, and Pan had the lowest ROM for ulnar deviation. These results can be attributed to differences in locomotor behaviour, as gibbons need a large wrist mobility in all directions for their highly arboreal lifestyle, whereas Macaca and Pan need a stable wrist during terrestrial locomotion. However, we found no correlation between distal ulna/triquetrum/hamate shape and maximal ROM during ulnar deviation and supination in the different primate taxa. A larger dataset, in combination with behavioural and biomechanical studies, is needed to establish form-function relationships of the primate hand, which will aid the functional interpretation of primate fossil remains.

High frequency of dental caries and calculus in dentitions from a British medieval town.

OBJECTIVE: Dental pathology and tooth wear data can offer valuable insights into the diet and behaviour of past populations. This study aimed to investigate the presence of dietary continuity by examining different types of dental pathology and tooth wear in a medieval sample from the United Kingdom, comparing them to earlier and later samples from the same location. DESIGN: A comprehensive examination was conducted on 41 individuals (comprising 914 permanent teeth) retrieved from the medieval cemetery of St. Owens Church in Southgate Street, Gloucester, UK. The research focused on documenting and analysing various types of dental pathology and tooth wear, such as dental caries, calculus, and tooth chipping. The frequency of these specific pathologies and wear patterns was then compared to existing literature. Additionally, non-masticatory tooth wear was also evaluated as part of the study. RESULTS: The sample exhibits high levels of carious lesions and calculus (24 % and 74 % of teeth respectively). Anterior teeth also show an elevated chipping frequency, and along with occlusal notches on the maxillary central incisors suggest teeth were regularly used for non-masticatory purposes. CONCLUSIONS: Caries frequency is similar to sites from later periods and may relate to the early adoption of consuming refined carbohydrates. However, remains from the same area, but the earlier Roman period, also shows high rates of caries and calculus, suggesting a continuation of consuming certain cariogenic foods, or certain behavioural/environmental factors, may instead be responsible for these pathology and wear patterns.

MEMOR: A database of archeological human remains collections from Flanders, Belgium.

The aim of this article is to describe a newly created open access database of archeological human remains collections from Flanders, Belgium. The MEMOR database (www.memor.be) was created to provide an overview of the current practices of loans, reburial, and the research potential of human skeletons from archeological sites currently stored in Flanders. In addition, the project aimed to provide a legal and ethical framework for the handling of human remains and was created around stakeholder involvement from anthropologists, geneticists, contract archeologists, the local, regional and national government agencies, local and national government, universities, and representatives of the major religions. The project has resulted in the creation of a rich database with many collections available for study. The database was created using the open-source Arches data management platform that is freely available for organizations worldwide to configure in accordance with their individual needs and without restrictions on its use. Each collection is linked to information about the excavation and the site the remains originate from, its size and time period. In addition, a research potential tab reveals whether any analyses were performed, and whether excavation notes are available with the assemblage. The database currently contains 742 collections, ranging in size from 1 to over 1000 individuals. New collections will continue to be added when new assemblages are excavated and studied. The database can also be expanded to include human remains collections from other regions and other material categories, such as archaeozoological collections.

First insights into human mobility in Neolithic Belgium using strontium isotopic analysis and proteomics: A case study of Grotte de La Faucille (Sclayn, province of Namur).

OBJECTIVES: So far, no 87 Sr/86 Sr mobility studies have been done for Neolithic remains from Belgium and information on the Sr isotopic variability in the region is scarce. This study aims to explore mobility in a Final Neolithic population from the funerary cave 'Grotte de La Faucille', contribute to the understanding of the isotopic composition of bioavailable Sr in Belgium, assess evidence for male mobility using proteomic analysis, and explore possible places of origin for nonlocal individuals. MATERIALS AND METHODS: The 87 Sr/86 Sr isotope ratio of dental enamel from six adults and six juveniles was determined. Liquid chromatography mass spectrometry-based protein analysis was employed to identify individuals of male biological sex. 87 Sr/86 Sr of micromammal teeth, snail shells, and modern plants from three geological areas in Belgium were measured to establish isotopic signatures for bioavailable strontium. Nonlocality was assessed by comparing human 87 Sr/86 Sr isotope ratios to the 87 Sr/86 Sr range for bioavailable Sr. RESULTS: Four individuals yielded 87 Sr/86 Sr isotope ratios consistent with a nonlocal origin. No statistical differences were found between adults and juveniles. Three males were detected in the sample set, of which two show nonlocal 87 Sr/86 Sr values. DISCUSSION: This study provides evidence for mobility in Final Neolithic Belgium. The four nonlocal 87 Sr/86 Sr signatures correspond with the 87 Sr/86 Sr of bio-available Sr in Dutch South Limburg, the Black Forest in Southwest Germany, and regions of France, such as parts of the Paris Basin and the Vosges. The results support the ruling hypothesis of connections with Northern France, brought to light by archeological research.

Reconstructing Articular Cartilage in the Australopithecus afarensis Hip Joint and the Need for Modeling Six Degrees of Freedom.

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.

One size fits all? Stature estimation from footprints and the effect of substrate and speed on footprint creation.

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.

Functional signals and covariation in triquetrum and hamate shape of extant primates using 3D geometric morphometrics.

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.

Reevaluating the timing of Neanderthal disappearance in Northwest Europe.

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.

Locomotion, posture, and the foramen magnum in primates: Reliability of indices and insights into hominin bipedalism.

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.

The morphological affinity of the Early Pleistocene footprints from Happisburgh, England, with other footprints of Pliocene, Pleistocene, and Holocene age.

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.

Sex differences in the patterning of age-related bone loss in the human hallucal metatarsal in rural and urban populations.

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.

The functional significance of dental and mandibular reduction in Homo: A catarrhine perspective.

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.

Root grooves on two adjacent anterior teeth of Australopithecus africanus.

Tooth root grooves and other ante-mortem dental tissue loss, not associated with caries found on or near the cementoenamel junction (CEJ), are commonly termed non-carious cervical lesions. Three main processes are implicated in forming these lesions: abrasion, dental erosion, and abfraction. As yet, these lesions have not been described in non-Homo hominins. In this study, South African fossil hominin collections were examined for evidence of any type of non-carious cervical lesion. Only one individual shows ante-mortem root grooves consistent with non-carious cervical lesions. Two teeth, a mandibular right permanent lateral incisor (STW 270) and canine (STW 213), belonging to the same Australopithecus africanus individual, show clear ante-mortem grooves on the labial root surface. These lesions start below the CEJ, extend over a third of the way toward the apex, and taper to a point towards the lingual side. The characteristics of these grooves suggest the predominant aetiology was erosive wear. In addition, they are extremely similar to clinical examples of dental erosion. These are the oldest hominin examples of non-carious cervical lesions and the first described in a genus other than Homo. Further, the lesions suggest that this individual regularly processed and consumed acidic food items.

Neuromandibular integration in humans and chimpanzees: Implications for dental and mandibular reduction in Homo.

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.

Amelogenesis imperfecta in the dentition of a wild chimpanzee.

This report describes a case of amelogenesis imperfecta in the dentition of a female chimpanzee. Amelogenesis imperfecta is a group of rare genetic conditions that create severe enamel defects, which, although well researched in humans, has not yet been investigated in wild non-human primates.

Behavioral inferences from the high levels of dental chipping in Homo naledi.

OBJECTIVES: 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. 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 and in samples of nonhuman primates (n = 3) and recent humans (n = 7). RESULTS: 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%). 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 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.