Geometric morphometric variability in the supraorbital and orbital region of Middle Pleistocene hominins: Implications for the taxonomy and evolution of later Homo.

This study assessed variation in the supraorbital and orbital region of the Middle Pleistocene hominins (MPHs), sometimes called Homo heidelbergensis s.l., to test whether it matched the expectations of intraspecific variation. The morphological distinctiveness and relative variation of this region, which is relatively well represented in the hominin fossil record, was analyzed quantitatively in a comparative taxonomic framework. Coordinates of 230 3D landmarks (20) and sliding semilandmarks (210) were collected from 704 specimens from species of Homo, Australopithecus, Paranthropus, Gorilla, Pan, Papio, and Macaca. Results showed that the MPHs had expected levels of morphological distinctiveness and intragroup and intergroup variation in supraorbital and orbital morphology, relative to commonly recognized non-hominin catarrhine species. However, the Procrustes distances between this group and H. sapiens were significantly higher than expected for two closely related catarrhine species. Furthermore, this study showed that variation within the MPH could be similarly well contained within existing hypodigms of H. sapiens, H. neanderthalensis, and H. erectus s.l. Although quantitative assessment of supraorbital and orbital morphology did not allow differentiation between taxonomic hypotheses in later Homo, it could be used to test individual taxonomic affiliation and identify potentially anomalous individuals. This study confirmed a complicated pattern of supraorbital and orbital morphology in the MPH fossil record and raises further questions over our understanding of the speciation of H. sapiens and H. neanderthalensis and taxonomic diversity in later Homo.

Twenty-first century bioarchaeology: Taking stock and moving forward.

This article presents outcomes from a Workshop entitled "Bioarchaeology: Taking Stock and Moving Forward," which was held at Arizona State University (ASU) on March 6-8, 2020. Funded by the National Science Foundation (NSF), the School of Human Evolution and Social Change (ASU), and the Center for Bioarchaeological Research (CBR, ASU), the Workshop's overall goal was to explore reasons why research proposals submitted by bioarchaeologists, both graduate students and established scholars, fared disproportionately poorly within recent NSF Anthropology Program competitions and to offer advice for increasing success. Therefore, this Workshop comprised 43 international scholars and four advanced graduate students with a history of successful grant acquisition, primarily from the United States. Ultimately, we focused on two related aims: (1) best practices for improving research designs and training and (2) evaluating topics of contemporary significance that reverberate through history and beyond as promising trajectories for bioarchaeological research. Among the former were contextual grounding, research question/hypothesis generation, statistical procedures appropriate for small samples and mixed qualitative/quantitative data, the salience of Bayesian methods, and training program content. Topical foci included ethics, social inequality, identity (including intersectionality), climate change, migration, violence, epidemic disease, adaptability/plasticity, the osteological paradox, and the developmental origins of health and disease. Given the profound changes required globally to address decolonization in the 21st century, this concern also entered many formal and informal discussions.

A dental revolution: The association between occlusion and chewing behaviour.

Dentistry is confronted with the functional and aesthetic consequences that result from an increased prevalence of misaligned and discrepant dental occlusal relations in modern industrialised societies. Previous studies have indicated that a reduction in jaw size in response to softer and more heavily processed foods during and following the Industrial Revolution (1,700 CE to present) was an important factor in increased levels of poor dental occlusion. The functional demands placed on the masticatory system play a crucial role in jaw ontogenetic development; however, the way in which chewing behaviours changed in response to the consumption of softer foods during this period remains poorly understood. Here we show that eating more heavily processed food has radically transformed occlusal power stroke kinematics. Results of virtual 3D analysis of the dental macrowear patterns of molars in 104 individuals dating to the Industrial Revolution (1,700-1,900 CE), and 130 of their medieval and early post-medieval antecedents (1,100-1,700 CE) revealed changes in masticatory behaviour that occurred during the early stages of the transition towards eating more heavily processed foods. The industrial-era groups examined chewed with a reduced transverse component of jaw movement. These results show a diminished sequence of occlusal contacts indicating that a dental revolution has taken place in modern times, involving a dramatic shift in the way in which teeth occlude and wear during mastication. Molar macrowear suggests a close connection between progressive changes in chewing since the industrialization of food production and an increase in the prevalence of poor dental occlusion in modern societies.

A critical assessment of the potential for Structure-from-Motion photogrammetry to produce high fidelity 3D dental models.

OBJECTIVES: The recent proliferation of methods of 3D model generation has enabled the development of new approaches to the analysis of dental form, function and wear. This article assesses whether Structure-from-motion (SfM) photogrammetry is capable of producing virtual 3D models of teeth of adequate quality for assessing fine scale surface details, such as dental macrowear patterns. Reference models were generated using a high resolution structured light scanner to assess the accuracy of the photogrammetric models generated. MATERIALS AND METHODS: Dental gypsum models of the molar teeth of human individuals from St. Michael's Litten, Chichester, Post-medieval assemblage (n = 17) were used for 3D model generation. Photogrammetry was performed using Agisoft Metashape and reference 3D models were generated using a GOM ATOS 80 scanner. Focus stacking was explored as a method of enhancing 3D model detail. Differences between the photogrammetric and reference models were assessed using CloudCompare and the quality of the surface detail was examined quantitatively using Occlusal Fingerprint Analysis. RESULTS: Photogrammetric model generation was highly replicable and the tooth models produced closely approximated the overall geometry of those derived from the structured light scanner. Dental wear facet area measurements on the photogrammetric models differed significantly, however, from those derived from the structured light scanning reference models. DISCUSSION: Photogrammetry can create virtual dental models from which crude quantitative size and shape data can be obtained. Finer scale surface details are not accurately reproduced on SfM models using the methods outlined in the current article due to high levels of surface noise.

Taxonomic variation in the supraorbital region of catarrhine primates.

OBJECTIVES: This study aimed to test the taxonomic utility of the catarrhine supraorbital region using 3D geometric morphometrics, with the aim of establishing its potential use in elucidating the position of more debated hominin groups. MATERIALS AND METHODS: 230 3D coordinates were used to record the supraorbital morphology of two datasets: one containing 460 non-hominin catarrhine primates from species and subspecies of Gorilla, Pan, Papio, and Macaca; and the other containing 55 Pleistocene hominins from Homo, Australopithecus, and Paranthropus. Principal component analyses in tangent, form, and allometry-free shape space were used to assess differentiation of taxa, with biological distinctiveness of taxa being established using step-wise discriminant analysis with subsampling. RESULTS: Results indicated that the recorded supraorbital morphology could be used to separate non-hominin catarrhine primate genera, species, and subspecies, although accuracy was found to decrease with decreasing Linnaean rank. In addition, analyses in tangent space were found to produce the highest accuracy when classifying primates of known taxonomy. Biological distinctiveness of the middle and later Homo species was comparable to or higher than that of the non-hominin primates, and relatively lower for the earlier groups of Homo. DISCUSSION: This study indicates that the supraorbital region preserves taxonomic information that can be used to delineate between closely related groups, both within hominins and wider catarrhine primates. Therefore, this region may be used to provide insight when assessing the taxonomic affiliation of disputed hominin specimens.

Nondestructive adult age at death estimation: Visualizing cementum annulations in a known age historical human assemblage using synchrotron X-ray microtomography.

OBJECTIVES: Adult age at death estimation continues to challenge physical anthropologists. One estimation method involves counting tooth cementum annulations (TCA). Non-destructively accessing TCA is a critical step to approaching fossil teeth of unknown age and to verifying life history profiles of human ancestors. This pilot study aims to (a) non-destructively image TCA in teeth from a known age archeological human population by propagation phase contrast X-ray synchrotron µCT (PPC-SR-µCT) (b) test the correlation between real and estimated ages, and the accuracy, precision and bias of age estimates. MATERIALS AND METHODS: We examine 20 permanent human canines (aged 20-81 years), from a 18th to 19th century known age collection from St. Luke's Church (London, England). We scanned transverse segments of acellular cementum in the apical portion of the middle root third using PPC-SR-µCT. We generated virtual transverse sections on which two observers perform two sessions of blind TCA counts. We calculate the estimated ages at death by adding 10 years to the TCA counts. RESULTS: A moderately strong positive linear relationship exists between real and estimated ages (r = 0.76, p < .001), with an average inaccuracy of 16.1 years and an average bias towards underestimation of 15.7 years. This difference is lower in individuals <50 years (6.8 and 6.5 years, respectively, n = 10) compared with those >50 years (24.9 years, n = 10). DISCUSSION: We reliably imaged and identified TCA in individuals <50 years from a known-age archeological sample. Scanning refinement will yield a promising alternative to current destructive methods of TCA analyses and to aid access to life history events in adult fossil hominins.

Application of the Bang and Ramm age at death estimation method to two known-age archaeological assemblages.

The Bang and Ramm method uses root dentine translucency (RDT) length in sectioned or unsectioned teeth as a sole indicator of chronological age at death in adult human remains. The formulae have been tested on modern remains of known age and on modern and archaeological remains of unknown age. This is the first published study of the method on known-age archaeological specimens and tests whether RDT is a good indicator of chronological age in buried human remains. We applied the Bang and Ramm equations to two 18th and 19th century assemblages excavated from the crypt of Christ Church, Spitalfields, and the cemetery of All Hallows by the Tower. Translucency was defined by shining a light through the external unsectioned root surface and was measured from digital images of 583 and 83 nonmolar roots from 126 Spitalfields and 12 All Hallows individuals, respectively, aged 21-81 years. Average absolute difference between real age and estimated age was 10.7 years and 8.4 years for Spitalfields and All Hallows individuals, respectively, with 58% and 75% estimated within 10 years of known age, and 29% and 33% estimated within five years of known age. These estimations are comparable to results from other ageing methods applied to the Spitalfields collection. Ages from both populations were estimated largely to the middle ranges, with younger individuals overestimated and older individuals underestimated. This is a common occurrence when using inverse calibration, where age is treated as the dependent variable and the ageing feature as the independent variable.

Changes in mandibular dimensions during the mediaeval to post-mediaeval transition in London: a possible response to decreased masticatory load.

OBJECTIVES: Biomechanical forces, such as those produced during mastication, are considered a primary agent in stimulating craniofacial growth and development. There appears to be a strong connection between the strength of the masticatory muscles and the dimensions of the craniofacial complex, with changes in biomechanical force and muscular strength influencing and altering the underlying bony tissues. This is markedly apparent in the mandible and it is possible to infer that changes to mandibular form are due in part to dietary changes. This study aims to investigate this idea by using an archaeological sample from a period that experienced important dietary changes as a result of the Industrial Revolution. DESIGN: 279 skeletons from the mediaeval and post-mediaeval periods in London were selected for analysis, and a detailed metric examination of each mandible was carried out. RESULTS: Males and females were analysed separately and statistically significant reductions were observed in nearly all post-mediaeval measurements. This effect was most pronounced in the areas of the mandible associated with masticatory muscles attachment, including the gonial angle, ramus height and width, bi-gonial breadth and bi-condylar breadth. CONCLUSIONS: These recorded changes in mandibular morphology of mediaeval and post-mediaeval Londoners are most likely the result of a shift in diet (and associated decrease in masticatory function) observed in the period surrounding the Industrial Revolution.

Intrapopulation variation in macro tooth wear patterns--a case study from Igloolik, Canada.

The pattern of human tooth wear-the way it varies between teeth in the mouth-is crucial to our understanding of important questions in archeology and paleoanthropology, such as the contrasts in diet and behavior between Neanderthals and early modern humans in Europe and Asia, or with the adoption of agriculture in the Americas. Little is known, however, about the way in which wear patterns develop with increasing age or the way in which they differ between males and females. One explanation is that few living people show the high rates of tooth wear seen worldwide throughout the preindustrial archaeological record. The study described here investigates the macroscopic pattern of tooth wear in a unique group of known age and sex dental casts from living Canadian Inuit from Igloolik. The results show that the Igloolik people possessed a pattern of extremely heavy anterior tooth wear, relative to the first molar and the other posterior teeth, which is attributed to the use of the anterior teeth in cultural practices as well as the extreme and marginal environments in which they lived. Heavy anterior tooth wear was established at an early age and maintained throughout life; statistically significant differences were found between the wear patterns of males and females and are explained in terms of sexual division of labor within the community. This study highlights the need to understand both intra- and interpopulation variation in tooth wear patterns when interpreting patterns in past human groups.

Brief communication: the use of alternative dental measurements on deciduous teeth.

The present study investigates the utility of cervical measurements in deciduous teeth and how they correlate with traditional measurements of the crown. First, this study establishes definitions by which these measurements could reliably be taken. Next, deciduous cervical and traditional crown diameters were taken on three distinct skeletal samples: a Neolithic assemblage from Central Anatolia (Çatalhöyük, n = 85), a precontact sample from Northern California (CA-ALA-329, n = 34), and a group of intrusive burials interred at Çatalhöyük that date between AD 60 and 1650 (n = 38). Across the dentition there are positive correlations between crown and cervical measurements, which tend to be higher in anterior teeth than in posterior teeth. Both measurements show low correlations with age; however, cervical measurements show fewer negative correlations with age. An intraobserver error study found low levels of error for both types of measurements. On a subset of the Çatalhöyük sample (n = 9), a principal components and biological distance analysis were conducted comparing the two types of measurements. Also, all three samples were subject to a canonical discriminant function analysis and the results from cervical and crown measurements were compared. All analyses produced slightly different results for each type of data suggesting that crown and cervical measurements capture different aspects of tooth shape. While cervical and crown measurements provide different statistical results, cervical measurements can provide information relevant to anthropological studies and may allow for larger datasets to be used by allowing the inclusion of teeth with modified crowns.

Tooth wear, Neanderthal facial morphology and the anterior dental loading hypothesis.

The Anterior Dental Loading Hypothesis states that the unique Neanderthal facial and dental anatomy was an adaptive response to the regular application of heavy forces resulting from both the masticatory and cultural use of the anterior teeth. Heavy anterior tooth wear frequently observed in Neanderthal specimens is cited as a main source of evidence for heavy forces being applied to these teeth. From this, it might be predicted that the wear shown on the anterior teeth of Neanderthals would greatly exceed that of the posterior teeth and that this differential would be greater than in other hominins with different facial morphologies. In this paper, a new method of examining tooth wear patterns is used to test these predictions in a large assemblage of Late Pleistocene hominins and a group of recent hunter-gatherers from Igloolik, Canada. The results show that all Late Pleistocene hominins, including Neanderthals, had heavily worn anterior teeth relative to their posterior teeth but, contrary to expectations, this was more pronounced in the modern humans than in the Neanderthals. The Igloolik Inuit showed heavier anterior tooth wear relative to their posterior teeth than any Late Pleistocene hominins. There was, however, a characteristic Neanderthal pattern in which wear was more evenly spread between anterior teeth than in modern humans. Overall, the evidence presented here suggests that all Late Pleistocene hominins habitually applied heavy forces between their anterior teeth and that Neanderthals were not exceptional in this regard. These results therefore does not support the Anterior Dental Loading Hypothesis.

The earliest evidence for anatomically modern humans in northwestern Europe.

The earliest anatomically modern humans in Europe are thought to have appeared around 43,000-42,000 calendar years before present (43-42 kyr cal BP), by association with Aurignacian sites and lithic assemblages assumed to have been made by modern humans rather than by Neanderthals. However, the actual physical evidence for modern humans is extremely rare, and direct dates reach no farther back than about 41-39 kyr cal BP, leaving a gap. Here we show, using stratigraphic, chronological and archaeological data, that a fragment of human maxilla from the Kent's Cavern site, UK, dates to the earlier period. The maxilla (KC4), which was excavated in 1927, was initially diagnosed as Upper Palaeolithic modern human. In 1989, it was directly radiocarbon dated by accelerator mass spectrometry to 36.4-34.7 kyr cal BP. Using a Bayesian analysis of new ultrafiltered bone collagen dates in an ordered stratigraphic sequence at the site, we show that this date is a considerable underestimate. Instead, KC4 dates to 44.2-41.5 kyr cal BP. This makes it older than any other equivalently dated modern human specimen and directly contemporary with the latest European Neanderthals, thus making its taxonomic attribution crucial. We also show that in 13 dental traits KC4 possesses modern human rather than Neanderthal characteristics; three other traits show Neanderthal affinities and a further seven are ambiguous. KC4 therefore represents the oldest known anatomically modern human fossil in northwestern Europe, fills a key gap between the earliest dated Aurignacian remains and the earliest human skeletal remains, and demonstrates the wide and rapid dispersal of early modern humans across Europe more than 40 kyr ago.

Periodontal disease in a Portuguese identified skeletal sample from the late nineteenth and early twentieth centuries.

Periodontal status was investigated in 600 adult dentitions belonging to the identified osteological collections curated at the University of Coimbra, Portugal. These collections date to a point temporally intermediate between the large epidemiological studies of the 20th century and archaeological collections that antedate the 19th century. The aim of this study is to compare periodontal data derived from contemporary samples with statistics compiled from epidemiological studies to determine if factors such as age-at-death, sex, and tooth type are essential or not to account for in future archaeological studies of periodontitis. Periodontal disease status was assessed based on the textural and architectural variations of the interdental septum and the extent of bone loss. Overall, the frequency of periodontitis within the Coimbra collections is 73.8%. Men were more susceptible to periodontal disease than women. Gingivitis was widespread in the younger age groups. Destructive periodontitis was observed early in adulthood, rising steadily with age. The most susceptible sites to periodontal breakdown were located in the posterior region of the upper jaw. Some variation in reported frequencies of periodontitis was found in epidemiological reports, which might result from variation in methods for identifying the pathology, differences in the age composition of the samples examined, variation in oral hygiene and/or diet, or some other factors. Regarding the pattern of distribution of periodontal disease, Coimbra results were similar to comparable modern epidemiological surveys, making clear the importance of considering sex, age, and oral distribution of periodontitis in future archaeological studies.

Deciduous tooth growth in an ancient Greek infant cemetery.

The Kylindra cemetery on Astypalaia in the Dodecanese, in use 750 BC to 1st century AD, contains a unique skeletal collection. Over 2,400 infant inhumations, each buried in its own clay pot, have been uncovered so far. The skeletal material from each burial is embedded within a ball of accreted earth and since 2001, some 850 infant remains have been recovered and conserved. Most of these died perinatally, but some were very premature babies and some appear to have survived for several months after birth. A study to estimate ages at death of 277 teeth from 107 infants, using microstructural growth markers, is currently underway. One immediate objective is to help resolve the enigma of why such an unusually large number of infants were interred on such a remote Aegean island. Longer term objectives are to reconstruct the sequences of development of the different deciduous tooth types, providing new standards of growth for long bones, the skull and the dentition. This paper presents an interim report on the findings from the histological study, which has analysed 68 teeth from 36 individuals so far. Five of the 36 infants survived birth, three dying within the first 3 weeks of life and the other two surviving for about 3 months. Average appositional growth rates were found to be 3.6 microm/day, and initial mineralisation was found to be well below the figures in Sunderland and coworkers' study in 1987.

Dental caries in a Portuguese identified skeletal sample from the late 19th and early 20th centuries.

Dental caries was investigated in 600 adult dentitions belonging to the identified osteological collections of the Museum of Anthropology, University of Coimbra, Portugal (late 19th/early 20th centuries). The main advantage of this sample compared to an archaeological source is the presence of known demographic parameters such as age, sex, and occupation. The aim of this study is to investigate the issues involved in comparing caries data derived from archaeological death assemblages with statistics compiled from clinical studies of the living. When only the upper dentition was considered, higher rates were observed in females than in males. No differences were found between sexes for lower teeth. In both sexes, both the percentage of carious teeth and the severity of lesions were found to increase with age, demonstrating that caries activity continued throughout life. The slight decrease observed for the age group 70-79 years is probably due to the increased antemortem tooth loss in the elderly. Caries was most common at contact areas (32.9%) and rarest at smooth crown surfaces (6.5%). Root surface caries was graphed in relation to the exposure of roots, and it was confirmed that the degree of root exposure was not strongly related to the frequency of carious lesions on the exposed root surface, although both increased with age. Molars were attacked more frequently by caries as a whole than premolars, canines or incisors. The results are similar to studies of recent living populations with a limited access to professional dental care.

The developmental clock of dental enamel: a test for the periodicity of prism cross-striations in modern humans and an evaluation of the most likely sources of error in histological studies of this kind.

Dental tissues contain regular microscopic structures believed to result from periodic variations in the secretion of matrix by enamel- and dentine-forming cells. Counts of these structures are an important tool for reconstructing the chronology of dental development in both modern and fossil hominids. Most studies rely on the periodicity of the regular cross-banding that occurs along the long axis of enamel prisms. These prism cross-striations are widely thought to reflect a circadian rhythm of enamel matrix secretion and are generally regarded as representing daily increments of tissue. Previously, some researchers have argued against the circadian periodicity of these structures and questioned their use in reconstructing dental development. Here we tested the periodicity of enamel cross-striations--and the accuracy to which they can be used--in the developing permanent dentition of five children, excavated from a 19th century crypt in London, whose age-at-death was independently known. The interruption of crown formation by death was used to calibrate cross-striation counts. All five individuals produced counts that were strongly consistent with those expected from the independently known ages, taking into account the position of the neonatal line and factors of preservation. These results confirm that cross-striations do indeed reflect a circadian rhythm in enamel matrix secretion. They further validate their use in reconstructing dental development and in determining the age-at-death of the remains of children whose dentitions are still forming at the time of death. Significantly they identify the most likely source of error and the common difficulties encountered in histological studies of this kind.

Alternative dental measurements: proposals and relationships with other measurements.

Most archaeological and fossil teeth are heavily worn, and this greatly limits the usefulness of tooth crown diameter measurements, as they are usually defined at the widest points of the crown. There are alternatives, particularly measurements at the cervix of the tooth, where the crown joints the root, and measurements along a diagonal axis in molars, that are much less affected by wear. These would allow a wider range of specimens to be included, e.g., in the study of dental reduction in Upper Palaeolithic and Mesolithic Homo sapiens. In addition, they would allow the little-worn teeth of children to be compared directly with well-worn teeth in adults. These alternatives, however, have been little used, and as yet there have not been any studies of the repeatability with which they can be measured, or of the extent to which they are related to the more usual crown diameters. The present study is based on a group of unworn teeth, where direct comparisons could be made between the alternative measurements, which are not much affected by wear, with the usual crown diameters, which are very much affected. In an interobserver-error study of this material, cervical and diagonal measurements could be recorded as reliably as the usual crown diameters. The buccolingual cervical measurement was strongly correlated with the normal bucclingual crown diameter in all teeth, whereas the mesiodistal cervical measurement was highly correlated with the normal mesiodistal crown diameter in incisors and canines, but less so in premolars and molars. The molar diagonal measurements showed high correlations with all other measurements. Crown areas (robustness index) calculated from the usual diameters were strongly correlated with crown areas calculated from cervical measurements, and crown areas calculated from molar diagonals were strongly correlated with both other areas. Despite the long usage of the more usual maximum crown diameters, the alternative dental measurements could be measured just as reliably, could record similar information about tooth crown size, and would be better measures for the worn dentitions seen in archaeological and fossil material.