From hunter-gatherers to food producers: New dental insights into the Nile Valley population history (Late Paleolithic-Neolithic).

OBJECTIVES: This study presents biological affinities between the last hunter-fisher-gatherers and first food-producing societies from the Nile Valley. We investigate odontometric and dental tissue proportion changes between these populations from the Middle Nile Valley and acknowledge the biological processes behind them. MATERIALS AND METHODS: Dental remains of 329 individuals from Nubia and Central Sudan that date from the Late Pleistocene to the mid-Holocene are studied. Using 3D imaging techniques, we investigated outer and inner metric aspects of upper central incisors, and first and second upper molars. RESULTS: Late Paleolithic and Mesolithic foragers display homogeneous crown dimensions, dental tissue proportions, and enamel thickness distribution. This contrasts with Neolithic trends for significant differences from earlier samples on inner and outer aspects. Finally, within the Neolithic sample differences are found between Nubian and Central Sudanese sites. DISCUSSION: Substantial dental variation appears to have occurred around 6000 bce in the Nile Valley, coinciding with the emergence of food-producing societies in the region. Archeological and biological records suggest little differences in dietary habits and dental health during this transition. Furthermore, the substantial variations identified here would have happened in an extremely short time, a few centuries at most. This does not support in situ diet-related adaptation. Rather, we suggest these data are consistent with some level of population discontinuity between the Mesolithic and Neolithic samples considered here. Complex settlement processes could also explain the differences between Nubia and Central Sudan, and with previous results based on nonmetric traits.

Agenesis of the permanent teeth in sub-Saharan Africans: Prevalence, patterns, interpretations.

OBJECTIVE: Dental agenesis data in modern and premodern sub-Saharan Africans are presented by region, West, Central, East, and South, and by sex. Beyond characterizing the anomaly, comparisons are made with other populations and future work is encouraged. The findings should be of use to dental clinicians and anthropologists. METHODS: Agenesis of the UI2, LI1, UP2, LP2, UM3, and LM3 was recorded in 52 discrete samples of mainly skeletal dentitions (n = 2162) from across the subcontinent. After dividing into temporal categories, regional pooling was effected for adequate sample sizes across the vast geographic area. Only adults were included to record M3 status. Analyses included 95% confidence intervals and chi-square comparisons by region and sex. RESULTS: Of 1668 modern individuals 2.3% have UI2-LP2 agenesis (CI 1.6-3.1%). Regional and sex differences are non-significant, though females are most affected. For M3s it is 7.0% (5.7-8.4%), with the Central region sample differing significantly from the East and South. Females again have greater prevalence, with the difference in the West significant. UI2-LP2 agenesis affects 0.6% of 494 premodern individuals (0.1-1.8%), while M3 agenesis is 8.5% (6.1-11.5%). None of these differences are significant. CONCLUSIONS: Rates are toward the low end of global ranges, including 0.0-12.6% for UI2-LP2 from case reports, and 5.3-56.0% for M3 agenesis. With exceptions, generally insignificant inter-region differences imply that rates reasonably represent sub-Saharan peoples overall. Results will be of interest to anthropologists, but those related to risk factors, patterning, and prevalence may assist clinicians in tailoring treatment, while informing patients how this anomaly differs by population ancestry.

Sex-biased sampling may influence Homo naledi tooth size variation.

A frequent source of debate in paleoanthropology concerns the taxonomic unity of fossil assemblages, with many hominin samples exhibiting elevated levels of variation that can be interpreted as indicating the presence of multiple species. By contrast, the large assemblage of hominin fossils from the Rising Star cave system, assigned to Homo naledi, exhibits a remarkably low degree of variation for most skeletal elements. Many factors can contribute to low sample variation, including genetic drift, strong natural selection, biased sex ratios, and sampling of closely related individuals. In this study, we tested for potential sex-biased sampling in the Rising Star dental sample. We compared coefficients of variation for the H. naledi teeth to those for eight extant hominoid samples. We used a resampling procedure that generated samples from the extant taxa that matched the sample size of the fossil sample for each possible Rising Star dental sex ratio. We found that variation at four H. naledi tooth positions-I2, M1, P4, M1-is so low that the possibility that one sex is represented by few or no individuals in the sample cannot be excluded. Additional evidence is needed to corroborate this inference, such as ancient DNA or enamel proteome data, and our study design does not address other potential factors that would account for low sample variation. Nevertheless, our results highlight the importance of considering the taphonomic history of a hominin assemblage and suggest that sex-biased sampling is a plausible explanation for the low level of phenotypic variation found in some aspects of the current H. naledi assemblage.

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.

Inferring human neutral genetic variation from craniodental phenotypes.

There is a growing consensus that global patterns of modern human cranial and dental variation are shaped largely by neutral evolutionary processes, suggesting that craniodental features can be used as reliable proxies for inferring population structure and history in bioarchaeological, forensic, and paleoanthropological contexts. However, there is disagreement on whether certain types of data preserve a neutral signature to a greater degree than others. Here, we address this unresolved question and systematically test the relative neutrality of four standard metric and nonmetric craniodental data types employing an extensive computational genotype-phenotype comparison across modern populations from around the world. Our computation draws on the largest existing data sets currently available, while accounting for geographically structured environmental variation, population sampling uncertainty, disparate numbers of phenotypic variables, and stochastic variation inherent to a neutral model of evolution. Our results reveal that the four data types differentially capture neutral genomic variation, with highest signals preserved in dental nonmetric and cranial metric data, followed by cranial nonmetric and dental metric data. Importantly, we demonstrate that combining all four data types together maximizes the neutral genetic signal compared with using them separately, even with a limited number of phenotypic variables. We hypothesize that this reflects a lower level of genetic integration through pleiotropy between, compared to within, the four data types, effectively forming four different modules associated with relatively independent sets of loci. Therefore, we recommend that future craniodental investigations adopt holistic combined data approaches, allowing for more robust inferences about underlying neutral genetic variation.

Descriptive catalog of Homo naledi dental remains from the 2013 to 2015 excavations of the Dinaledi Chamber, site U.W. 101, within the Rising Star cave system, South Africa.

More than 150 hominin teeth, dated to ∼330-241 thousand years ago, were recovered during the 2013-2015 excavations of the Dinaledi Chamber of the Rising Star cave system, South Africa. These fossils comprise the first large single-site sample of hominin teeth from the Middle Pleistocene of Africa. Though scattered remains attributable to Homo sapiens, or their possible lineal ancestors, are known from older and younger sites across the continent, the distinctive morphological feature set of the Dinaledi teeth supports the recognition of a novel hominin species, Homo naledi. This material provides evidence of African Homo lineage diversity that lasts until at least the Middle Pleistocene. Here, a catalog, anatomical descriptions, and details of preservation and taphonomic alteration are provided for the Dinaledi teeth. Where possible, provisional associations among teeth are also proposed. To facilitate future research, we also provide access to a catalog of surface files of the Rising Star jaws and teeth.

Imputed genomes and haplotype-based analyses of the Picts of early medieval Scotland reveal fine-scale relatedness between Iron Age, early medieval and the modern people of the UK.

There are longstanding questions about the origins and ancestry of the Picts of early medieval Scotland (ca. 300-900 CE), prompted in part by exotic medieval origin myths, their enigmatic symbols and inscriptions, and the meagre textual evidence. The Picts, first mentioned in the late 3rd century CE resisted the Romans and went on to form a powerful kingdom that ruled over a large territory in northern Britain. In the 9th and 10th centuries Gaelic language, culture and identity became dominant, transforming the Pictish realm into Alba, the precursor to the medieval kingdom of Scotland. To date, no comprehensive analysis of Pictish genomes has been published, and questions about their biological relationships to other cultural groups living in Britain remain unanswered. Here we present two high-quality Pictish genomes (2.4 and 16.5X coverage) from central and northern Scotland dated from the 5th-7th century which we impute and co-analyse with >8,300 previously published ancient and modern genomes. Using allele frequency and haplotype-based approaches, we can firmly place the genomes within the Iron Age gene pool in Britain and demonstrate regional biological affinity. We also demonstrate the presence of population structure within Pictish groups, with Orcadian Picts being genetically distinct from their mainland contemporaries. When investigating Identity-By-Descent (IBD) with present-day genomes, we observe broad affinities between the mainland Pictish genomes and the present-day people living in western Scotland, Wales, Northern Ireland and Northumbria, but less with the rest of England, the Orkney islands and eastern Scotland-where the political centres of Pictland were located. The pre-Viking Age Orcadian Picts evidence a high degree of IBD sharing across modern Scotland, Wales, Northern Ireland, and the Orkney islands, demonstrating substantial genetic continuity in Orkney for the last ~2,000 years. Analysis of mitochondrial DNA diversity at the Pictish cemetery of Lundin Links (n = 7) reveals absence of direct common female ancestors, with implications for broader social organisation. Overall, our study provides novel insights into the genetic affinities and population structure of the Picts and direct relationships between ancient and present-day groups of the UK.

Hyperdontia across sub-Saharan Africa: Prevalence, patterning, and implications.

OBJECTIVE: Hyperdontia data in modern and premodern sub-Saharan Africans are presented by region-West, Central, East, and South, and sex. Beyond describing the anomaly, comparisons are made with other world populations and future work is promoted. These findings may be useful to both dental clinicians and anthropologists. METHODS: Hyperdontia presence and patterning were recorded in 51 samples of skeletal dentitions and hardstone casts (n = 1916). Its infrequency prompted regional pooling after grouping by time. Only adults were included to record later forming fourth molars reportedly common in Africans. Quantitative analyses, including 95% confidence intervals, were conducted to characterize spatiotemporally sub-Saharan peoples. RESULTS: Forty-four of 1429 modern individuals (3.08%) exhibit hyperdontia (CI 2.24-4.13%). Regional variation is significant, particularly West-Central vs. East-South, between 6.8% and 1.5%. Four of 487 premodern individuals, 0.82%, have hyperdontia (0.22-2.10%), with minimal regional differences. Males are most affected, as reported by other researchers. Other similarities with non-African research are evident regarding isomere, antimere, and form, with one key exception-a proclivity for the posterior over anterior maxilla. CONCLUSIONS: 3.08% is toward the upper end of published world ranges, including an oft-cited 0.1-3.6 + %. However, the regional variation argues against a single prevalence to describe collectively the subcontinental inhabitants. This variation parallels past west to east and south migrations like other biocultural indicators. Thus, beyond a health concern to clinicians or interesting anomaly to anthropologists, hyperdontia may be useful in other studies. There are no equivalent premodern ranges for comparison. Similarity in patterning overall to previous findings suggests a shared, potentially ancient genetic component in expression.

Dental caries in wild primates: Interproximal cavities on anterior teeth.

Dental caries has been reported in a variety of primates, although it is still considered rare in wild populations. In this study, 11 catarrhine primate taxa (n = 339 individuals; 7946 teeth) were studied for the presence of caries. A differential diagnosis of lesions in interproximal regions of anterior teeth was undertaken, since they had been previously described as both carious and non-carious in origin. Each permanent tooth was examined macroscopically, with severity and position of lesions recorded. Two specimens were examined further, using micro-CT scans to assess demineralization. Differential diagnosis confirmed the cariogenic nature of interproximal cavities on anterior teeth (ICATs). Overall results show 3.3% of all teeth (i.e., anterior and posterior teeth combined) were carious (n = 262), with prevalence varying among species from 0% to >7% of teeth affected. Those with the highest prevalence of ICATs include Pan troglodytes verus (9.8% of anterior teeth), Gorilla gorilla gorilla (2.6%), Cercopithecus denti (22.4%), Presbytis femoralis (19.5%), and Cercopithecus mitis (18.3%). ICATs make up 87.9% of carious lesions on anterior teeth. These results likely reflect dietary and food processing differences among species, but also between the sexes (e.g., 9.3% of all female P. troglodytes verus teeth were carious vs. 1.8% in males). Processing cariogenic fruits and seeds with the anterior dentition (e.g., wadging) likely contributes to ICAT formation. Further research is needed in living primate populations to ascertain behavioral/dietary influences on caries occurrence. Given the presence of ICATs in frugivorous primates, their diagnosis in archaeological and paleontological specimens may shed light on diet and food processing behaviors in fossil primates.

Ancient anomalies: Twinned and supernumerary incisors in a medieval Nubian.

During the analysis of a skeletal assemblage from a medieval cemetery in Nubia (c. AD 500-1550), a young adult female with abnormally developed maxillary incisors was discovered. The possible causes of the two dental anomalies found in this individual and their archaeological context are discussed. The remains are from a medieval assemblage from the Fourth Cataract region of Nubia, which forms part of the Nubian collection curated at the British Museum. The left central incisor has a twinned crown with two root canals, and a supernumerary tooth is present on the right side between the central incisor and lateral incisors. Although two different dental anomalies are present, the bilateral expression suggests that the same biological mechanism could be responsible.

Relative tooth size, Bayesian inference, and Homo naledi.

OBJECTIVES: Size-corrected tooth crown measurements were used to estimate phenetic affinities among Homo naledi (~335-236 ka) and 11 other Plio-Pleistocene and recent species. To assess further their efficacy, and identify dental evolutionary trends, the data were then quantitatively coded for phylogenetic analyses. Results from both methods contribute additional characterization of H. naledi relative to other hominins. MATERIALS AND METHODS: After division by their geometric mean, scaled mesiodistal and buccolingual dimensions were used in tooth size apportionment analysis to compare H. naledi with Australopithecus africanus, A. afarensis, Paranthropus robustus, P. boisei, H. habilis, H. ergaster, H. erectus, H. heidelbergensis, H. neanderthalensis, H. sapiens, and Pan troglodytes. These data produce equivalently scaled samples unaffected by interspecific size differences. The data were then gap-weighted for Bayesian inference. RESULTS: Congruence in interspecific relationships is evident between methods, and with many inferred from earlier systematic studies. However, the present results place H. naledi as a sister taxon to H. habilis, based on a symplesiomorphic pattern of relative tooth size. In the preferred Bayesian phylogram, H. naledi is nested within a clade comprising all Homo species, but it shares some characteristics with australopiths and, particularly, early Homo. DISCUSSION: Phylogenetic analyses of relative tooth size yield information about evolutionary dental trends not previously reported in H. naledi and the other hominins. Moreover, with an appropriate model these data recovered plausible evolutionary relationships. Together, the findings support recent study suggesting H. naledi originated long before the geological date of the Dinaledi Chamber, from which the specimens under study were recovered.

The transition from hunting-gathering to agriculture in Nubia: dental evidence for and against selection, population continuity and discontinuity.

Some researchers posit population continuity between Late Palaeolithic hunter-gatherers of the late Pleistocene and Holocene agriculturalists from Lower (northern) Nubia, in northeast Africa. Substantial craniodental differences in these time-successive groups are suggested to result from in situ evolution. Specifically, these populations are considered a model example for subsistence-related selection worldwide in the transition to agriculture. Others question continuity, with findings indicating that the largely homogeneous Holocene populations differ significantly from late Pleistocene Lower Nubians. If the latter are representative of the local populace, post-Pleistocene discontinuity is implied. So who was ancestral to the Holocene agriculturalists? Dental morphological analyses of 18 samples (1075 individuals), including one dated to the 12th millennium BCE from Al Khiday, near the Upper Nubian border, may provide an answer. It is the first Late Palaeolithic sample (n = 55) recovered within the region in approximately 50 years. Using the Arizona State University Dental Anthropology System to record traits and multivariate statistics to estimate biological affinities, Al Khiday is comparable to several Holocene samples, yet also highly divergent from contemporaneous Lower Nubians. Thus, population continuity is indicated after all, but with late Pleistocene Upper-rather than Lower Nubians as originally suggested-assuming dental traits are adequate proxies for ancient DNA.

Rocker jaw: Global context for a Polynesian characteristic.

Our goal is to describe the global distribution of the "rocker jaw" variant in human populations. Rocker jaw refers to mandibles that lack the antegonial notch, making them unstable on a flat surface. Data were collected by C.G. Turner II on 9,207 individuals from Asia, Europe, the Pacific, and the Americas, and by J.D. Irish on 3,526 individuals from North and South Africa. With a focus on Polynesia, where the trait is most common, frequencies are presented for subdivisions of Oceania, Australasia, Eurasia, the Americas, and Africa. While the rocker jaw is a Polynesian characteristic, the trait is found throughout the world. Within major geographic regions, there are interesting contrasts, for example, (a) the similarity of Jomon and Ainu and their difference from modern Japanese; (b) Aleuts and Northwest Coast Indians are similar and both are distinct from the Inuit and other Native Americans; and (c) North and Sub-Saharan Africans show a regional difference that parallels genetic and dental distinctions. Skeletons in South America that exhibit the rocker jaw have been interpreted as Polynesian voyagers who ventured to the west coast of South America. The rarity of rocker jaw in South American natives supports this view. The rocker jaw can be attributed to the unique basicranium morphology and large upper facial height of Polynesians, which highlights the integrated growth of a functional module (i.e., mastication) of the craniofacial complex. The unusually high frequency of the trait in Polynesians is a product of both function and founder effect/genetic drift.

Do dental nonmetric traits actually work as proxies for neutral genomic data? Some answers from continental- and global-level analyses.

OBJECTIVES: Crown and root traits, like those in the Arizona State University Dental Anthropology System (ASUDAS), are seemingly useful as genetic proxies. However, recent studies report mixed results concerning their heritability, and ability to assess variation to the level of genomic data. The aim is to test further if such traits can approximate genetic relatedness, among continental and global samples. MATERIALS AND METHODS: First, for 12 African populations, Mantel correlations were calculated between mean measure of divergence (MMD) distances from up to 36 ASUDAS traits, and FST distances from >350,000 single nucleotide polymorphisms (SNPs) among matched dental and genetic samples. Second, among 32 global samples, MMD and FST distances were again compared. Correlations were also calculated between them and inter-sample geographic distances to further evaluate correspondence. RESULTS: A close ASUDAS/SNP association, based on MMD and F ST correlations, is evident, with r m -values between .72 globally and .84 in Africa. The same is true concerning their association with geographic distances, from .68 for a 36-trait African MMD to .77 for F ST globally; one exception is F ST and African geographic distances, r m = 0.49. Partial MMD/F ST correlations controlling for geographic distances are strong for Africa (.78) and moderate globally (.4). DISCUSSION: Relative to prior studies, MMD/F ST correlations imply greater dental and genetic correspondence; for studies allowing direct comparison, the present correlations are markedly stronger. The implication is that ASUDAS traits are reliable proxies for genetic data-a positive conclusion, meaning they can be used with or instead of genomic markers when the latter are unavailable.

Tooth transposition prevalence and type among sub-Saharan Africans.

OBJECTIVES: Although rare, tooth transposition-an exchange in location of two teeth-is a frequent topic of study. Clinical and, to a much lesser extent, dental anthropological research have focused predominantly on prevalence (0.03%-0.74% in several world populations) and case studies, albeit on a restricted spatiotemporal scale. Many regions have received little attention, including sub-Saharan Africa, while premodern data are few. Here, the aim is to supplement both fields of dental research by reviewing previous publications, and newly reporting transposition rates, types, and co-occurring abnormalities in time-successive samples across the subcontinent. METHODS: Dental data in 51 sub-Saharan samples (>2500 individuals) dating >10 000 BC to 20th century were recorded. Of these, 36 are of modern and 15 premodern age, comprising males and females ≥12-years of age. Transposition presence, quadrant, and type were tabulated, cases described, and prevalence presented. In the latter case, Poisson 95% confidence intervals were calculated to better discern true population rates at various geographic levels. RESULTS: Overall, six of 1886 modern individuals (0.32%) and one of premodern age evidence Mx.C.P1, an exchange of the maxillary canine and first premolar. Various associated dental abnormalities are also evident, including retained deciduous teeth, reduced permanent crowns, and agenesis. CONCLUSIONS: This study provides additional insight into the geographic distribution, features, and time depth of transposition, along with hints supporting a genetic etiology and, potentially, some indications of diachronic change from an initial Mx.C.P1 to several types more recently based on premodern evidence. It is of clinical concern today, but is not just a modern anomaly.

Root caries on a Paranthropus robustus third molar from Drimolen.

OBJECTIVES: Dental caries is often perceived as a modern human disease. However, their presence is documented in many early human groups, various nonhuman primates and, increasingly, our hominin ancestors and relatives. In this study, we describe an antemortem lesion on the root of a Paranthropus robustus third molar from Drimolen, South Africa, which likely represents another example of caries in fossil hominins. MATERIALS AND METHODS: The molar, DNH 40, is dated to 2.0-1.5 Ma and displays a lesion on the mesial root surface, extending from the cementoenamel junction 3 mm down toward the apex. The position and severity of the lesion was macroscopically recorded and micro-CT scanned to determine the extent of dentine involvement. RESULTS: A differential diagnosis indicates root caries, as the lesion is indistinguishable from clinical examples. Although necrotic in appearance, external tertiary dentine is evident on a micro CT scan. Gingival recession and/or continuous eruption of the tooth as a result of extensive occlusal wear would have occurred to facilitate caries formation. Therefore, the lesion is likely linked to relative old age of this individual. DISCUSSION: This new example increases the total number of carious lesions described in P. robustus teeth to 10, on occlusal, interproximal, and now, root surfaces. Beyond the consumption of caries-causing food, caries formation would have also required the presence of requisite intraoral cariogenic bacteria in this individual and the species. Of interest, the presence of tertiary dentine on the outward surface suggests the DNH 40 lesion may have been arrested, that is, no longer active, perhaps relating to a change in diet or oral microbiome just prior to the individual's death.