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Objectives Linear enamel hypoplasia (LEH) is a common skeletal marker of physiological stress (e.g., malnutrition or illness) that is studied within and across populations, without reference to familial risk. We examine LEH prevalence in a population with known genealogical relationships to determine the potential influence of genetic heritability and shared environment. Methods LEH data of 239 individuals from a single population were recorded from the Ohio State University Menegaz-Bock collection dental casts. All individuals were of known age, sex, and genealogy. Narrow-sense heritability estimates were obtained for LEH presence and count data from all unworn, fully erupted teeth (excluding third molars) using SOLAR (v.8.1.1). Age, sex, and age-sex interaction were included as covariates. Models were re-run with a household effect variable. Results LEH persists across generations in this study population with moderate, significant heritability estimates for presence in four teeth, and count in four teeth (three teeth were significant for both). When a household effect variable was added, no residual heritability remained for LEH count on any tooth. There was no significant household effect for three of the four teeth that had significant heritability estimates for LEH presence. Age was a significant covariate. Further analyses with birth year data revealed a secular trend toward less LEH. Conclusions This study provides evidence for familial risk of LEH (genetic and environmental) that has consequences for the broad use of this skeletal marker of stress. These results have repercussions for archaeological assemblages, or population health studies, where genetic relatives and household groups might be heavily represented.

OBJECTIVES This article presents estimates of narrow-sense heritability and bivariate genetic correlation for 14 tooth crown morphological variants scored on permanent premolars, first molars, and second molars. The objective is to inform data collection and analytical practices in dental biodistance and to provide insights on the development of molar crowns as integrated structures. MATERIALS AND METHODS African American dental casts from the Menegaz-Bock collection were recorded for the Arizona State University Dental Anthropology System. Estimates of narrow-sense heritability and genetic correlation were generated using SOLAR v.8.1.1, which included assessment of age, sex, and birth year as covariates. Both continuous scale and dichotomized estimates are provided. RESULTS Heritability estimates were nonsignificant for the majority of variables; however, for variables yielding significant estimates, values were moderate to high in magnitude and comparable to previous studies. Comparing left and right-side heritability estimates suggests directional asymmetry in the expression of environmental variance, something not seen in anterior tooth traits. Genetic correlations were moderate among antimeres and metameres and low for different traits scored on the same tooth crown. Although several negative correlations were noted, few reached statistical significance. Results affirm some of the current data cleaning and analytical practices in dental biodistance, but others are called into question. These include the pooling of males and females and combining left and right-side data into a single dataset. CONCLUSIONS In comparison to anterior tooth crown traits, postcanine heritabilities were more often non-significant; however, those traits with significant heritability also tended to produce higher estimates. Genetic correlations were unremarkable, in part, because they were underpowered. However, M1 results may provide insight into the complex relationship between genes, environment, and development in determining ultimate crown form.

Studies of enamel growth and thickness, whether in paleoanthropology, bioarchaeology, or primatology, require measurements of crown height (CH), cuspal enamel thickness (CET), average (AET), and/or regional enamel thickness (RegAET) on complete, unworn crowns. Yet because fully unworn crowns are uncommon, three methods to bolster sample sizes by reconstructing slightly worn teeth have been developed: Profile, Polynomial, and Pen Tool. Although these methods have been tested for accuracy, no study has yet directly compared the three methods to assess their performance across CH, CET, AET, and RegAET measurements. Moreover, it is currently unclear how accurate the methods are when reconstructing crowns with varying degrees of wear. The present study addresses this gap in our understanding of how these methods perform on four key dental measurements, evaluates the degree of wear for which accurate crown reconstructions can be completed, and offers recommendations for applying these methods. Here, the methods are compared on Paranthropus robustus mandibular molars, a sample chosen because it exhibits variable morphology, presenting a challenge for reconstruction methods. For minimally worn teeth, Profile, Polynomial, and Pen Tool methods can be employed (in that order) for all measurements except CET, which cannot be reliably measured on reconstructions. For teeth with wear that obliterates the nadir of the occlusal basin or dentin horns, CH and AET can be measured using Profile and Polynomial reconstructions; however, no other measurements or methods were reliable. Recommendations provided here will make it possible to increase sample sizes and replicability, enhancing studies of enamel thickness and growth.

Objectives: Human tooth enamel retains evidence of growth in the form of Retzius lines. The number of daily growth increments between the regularly occurring lines defines their repeat interval, or periodicity. Retzius periodicity is often incorporated into enamel formation times, age-at-death reconstructions, or used to provide a basis from which to explore an underlying biorhythm. Biological anthropologists typically assume that RP remains constant within an individual and does not vary along the tooth-row. Here, we test that assumption. Materials and methods: RP was calculated from n = 223 thin sections of human permanent teeth from individuals of British and southern African origin. Forty individuals provided multiple teeth (n = 102 teeth) and a further 121 individuals each provided a single tooth. Results: We report first evidence that RP of permanent teeth does not always remain constant within an individual. Of those individuals that provided multiple teeth, 42% (n = 17/40) demonstrated a decrease in RP along the tooth row, with most shifting by two or more days (n = 11). Across the entire sample, mean RP of anterior teeth was significantly higher than molars. Mean premolar RP tended to be intermediate between anterior teeth and molars.Discussion: Our data do not support the assumption that RP invariably remains constant within the permanent teeth of an individual. Transferring RP from molars to incisors within an individual can result in a miscalculation of formation time and age-at-death by up to one year. Implications for biological anthropologists and the source of the underlying long period biorhythm are discussed.

OBJECTIVE Linear enamel hypoplasia (LEH) is a condition marked by localized reductions in enamel thickness, resulting from growth disruptions during dental development. We use quantitative criteria to characterize the depth of LEH defects and "normal" perikymata in great apes. We test the hypothesis that mountain gorillas have shallow defects compared to other taxa, which may have led to their underestimation in previous studies. MATERIALS AND METHODS Previous attempts to characterize LEH morphology quantitatively have been limited in sample size and scope. We generated digital elevation models using optical profilometry (Sensofar PLu Neox) and extracted 2D coordinates using ImageJ to quantify depths in canines from three great ape genera (N = 75 perikymata; 255 defects). RESULTS All defect depths fall outside the distribution of perikymata depths. Mountain gorilla defects are significantly shallower than those of other great ape taxa examined, including western lowland gorillas. Females have significantly deeper defects than males in all taxa. The deepest defect belongs to a wild-captured zoo gorilla. Virunga mountain gorilla specimens collected by Dian Fossey exhibit deeper defects than those collected recently. DISCUSSION Shallow defect morphology in mountain gorillas may have led to an underestimation of LEH prevalence in past studies. Defect depth is used as a proxy for insult severity, but depth might be influenced by inter- and intra-specific variation in enamel growth. Future studies should test whether severe insults are associated with deeper defects, as might be the case with Haloko, a wild-captured gorilla. Ongoing histologic studies incorporating associated behavioral records will test possible factors that underlie differences in defect morphology.

Enamel formation front (EFF) angles represent the leading edge of enamel matrix secretion at particular points in time. These angles are influenced by rates of enamel extension (the rates at which tooth crowns grow in height), rates of enamel matrix secretion and the angles that prisms make with the enamel-dentine junction. Previous research suggests, but has not yet established, that these angles reflect aspects of primate biology related to their pace of growth and development, most notably brain and body size. The present study tested this possibility on histological sections using phylogenetically-controlled and Bonferroni-corrected analyses spanning a broad taxonomic range. Ten species were represented in the analysis of anterior teeth; 17 in the analysis of posterior (postcanine) teeth (with varying sample sizes). Also, tested was the relationship of EFF angles to striae of Retzius periodicity (long period growth rhythms in enamel) and degree of folivory, as both factors are related to primate developmental rates. Finally, several analyses were conducted to investigate whether tooth size (operationalized as EDJ length) might mediate these relationships. Central results are as follows: (1) Relationships between EFF angles and brain weight (anterior teeth) and between EFF angles and body mass (anterior and posterior teeth) are statistically significant and (2) Mid-crown EFF angles are not statistically significantly related to EDJ lengths. These results suggest that tooth size does not mediate relationships between EFF angles and brain weight/body mass and are discussed with respect to underlying enamel growth variables (especially rates of enamel extension and secretion). Anat Rec, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 301:125-139, 2018. © 2017 Wiley Periodicals, Inc.

Objectives This article provides estimates of narrow-sense heritability and genetic pleiotropy for mesiodistal tooth dimensions for a sample of 20th century African American individuals. Results inform biological distance analysis and offer insights into patterns of integration in the human dentition. Materials and Methods Maximum mesiodistal crown dimensions were measured using Hillson-FitzGerald calipers on 469 stone dental casts from the Menegaz-Bock Collection. Narrow-sense heritability estimates and genetic and phenotypic correlations were estimated using SOLAR 8.1.1 with covariate screening for age, sex, age*sex interaction, and birth year. Results Heritability estimates were moderate (∼0.10 – 0.90; h2 mean = 0.51) for most measured variables with sex as the only significant covariate. Patterns of genetic correlation indicate strong integration across tooth classes, except molars. Comparison of these results to previously published work suggests lower overall heritability relative to other human populations and much stronger genetic integration across tooth classes than obtained from nonhuman primate genetic pleiotropy estimates. Conclusions These results suggest that the high heritabilities previously published may reflect overestimates inherent in previous study designs; as such the standard estimate of 0.55 used in biodistance analyses may not be appropriate. For the Gullah, isolation and endogamy coupled with elevated levels of physiological and economic stress may suppress narrow-sense heritability estimates. Pleiotropy analyses suggest a more highly integrated dentition in humans than in other mammals.

Deeper or more ‘severe’ linear enamel hypoplasia (LEH) defects are hypothesized to reflect more severe stress during development, but it is not yet clear how depth is influenced by intrinsic enamel growth patterns. Recent work documented inter- and intraspecific differences in LEH defect depth in extant great apes, with mountain gorillas having shallower defects than other taxa, and females having deeper defects than males. Here, we assess the correspondence of inter- and intraspecific defect depth and intrinsic aspects of enamel growth: enamel extension rates, outer enamel striae of Retzius angles, and linear enamel thickness. Thin sections of great ape canines (n = 40) from Gorilla beringei beringei, Gorilla gorilla gorilla, Pan troglodytes, and Pongo spp. were analyzed. Enamel extension rates were calculated within deciles of enamel-dentine junction length. Linear enamel thickness and the angle of intersection between striae of Retzius and the outer enamel surface were measured in the imbricational enamel. Mountain gorillas have faster enamel extension rates and shallower striae angles than the other taxa examined. Mountain gorillas have thinner imbricational enamel than western lowland gorillas and orangutans, but not chimpanzees. In the combined-taxon sample, females exhibit larger striae angles and thicker imbricational enamel than males. Enamel extension rates are highly negatively correlated with striae angles and LEH defect depth. Enamel growth variation corresponds with documented inter- and intraspecific differences in LEH defect depth in great ape canines. Mountain gorillas have shallower striae angles and faster extension rates than other taxa, which might explain their shallow LEH defect morphology and the underestimation of their LEH prevalence in previous studies. These results suggest that stressors of similar magnitude and timing might produce defects of different depths in one species or sex vs. another, which has implications for interpretations of stress histories in hominins with variable enamel growth patterns.

Objectives To update histological data on crown initiation and completion ages in southern Africans. To evaluate implications of these data for studies that: (a) rely on these data to time linear enamel hypoplasias (LEHs), or, (b) use these data for comparison to fossil hominins. Materials and methods Initiation ages were calculated on 67 histological sections from southern Africans, with sample sizes ranging from one to 11 per tooth type. Crown completion ages for southern Africans were calculated in two ways. First, actual derived initiation ages were added to crown formation times for each histological section to obtain direct information on the crown completion ages of individuals. Second, average initiation ages from this study were added to average crown formation times of southern Africans from the Reid and coworkers previous studies that were based on larger samples. Results For earlier-initiating tooth types (all anterior teeth and first molars), there is little difference in ages of initiation and crown completion between this and previous studies. Differences increase as a function of initiation age, such that the greatest differences between this and previous studies for both initiation and crown completion ages are for the second and third molars. Discussion This study documents variation in initiation ages, particularly for later-initiating tooth types. It upholds the use of previously published histological aging charts for LEHs on anterior teeth. However, this study finds that ages of crown initiation and completion in second and third molars for this southern African sample are earlier than previously estimated. These earlier ages reduce differences between modern humans and fossil hominins for these developmental events in second and third molars.

Dental replicas are frequently utilized in paleoanthropological studies of perikymata and enamel hypoplasia. However, fossil teeth are often fragile and worn, causing two problems: (1) the risk of damage by removing enamel fragments when impression-making material is separated from the fossil tooth surface, and (2) the need to reconstruct worn portions of the crown to assess perikymata number, distribution, and hypoplasia timing. This study presents the advantages of μCT data of canines and lateral incisors for (1) detecting cracks along the enamel-dentine junction (EDJ) which could cause damage when casting, and (2) reliably and non-destructively reconstructing worn or broken cusps. Fragile teeth of Homo naledi, Miocene, and Pleistocene specimens were μCT-scanned: 2D virtual sections and 3D models allowed for inspecting crack pattern beyond the external surface and 2D virtual sections were used to digitally reconstruct cusp tips (only Homo naledi). Micro-CT scans allowed cracks running along the EDJ and communicating with radial cracks in the enamel to be identified prior to casting. Cusp reconstructions using μCT data were conducted as precisely as when using thin-sections or photographs, and with high intra- and inter-observer agreement, while preserving the original specimen and affording numerous planes of virtual section. When available, μCT data should be inspected prior to tooth casting to exclude teeth that show a pattern of cracks that could lead to damage. Virtual sections allow for accessible, reliable, and non-destructive cusp reconstructions that may be used for developmental (e.g., perikymata and enamel hypoplasia) or enamel thickness studies. Anat Rec, 302:1516-1535, 2019. © 2018 American Association for Anatomy.

Linear enamel hypoplasias (LEH) have been used in physical anthropology as stress indicators. While circumstances such as malnutrition and illness certainly disrupt enamel formation, intrinsic features of enamel growth may affect defect expression as well. Variation in enamel growth factors, particularly lateral enamel formation time and the angle that striae of Retzius make with the outer enamel surface, may influence the number and visibility of enamel defects on teeth of different species and sexes. Understanding how such intrinsic factors affect enamel defect expression is critical to interpreting enamel defect differences between hominin species and groups. Here, the number of LEH defects and rates of enamel defect acquisition of two sympatric cercopithecoids (Macaca fascicularis and Trachypithecus cristatus) are compared. Their enamel formation time and perikymata spacing are considered in relation to their expression of defects. Longer lateral enamel formation times are not significantly associated with higher enamel defect counts in these two species or between conspecific sexes. Perikymata spacing (which is sometimes correlated with striae of Retzius angles) does not help to explain species-level differences in number or rate of defect acquisition either. However, the greater number of defects and rate of acquisition in females of both species, coupled with tighter perikymata spacing, suggests a possible role of striae angles influencing defect perceptibility. Significantly more research regarding the role enamel growth variables play in enamel defect expression is required for LEH to be used as an informative stress indicator in bioarchaeology and paleoanthropology.

Objectives This article presents estimates of narrow-sense heritability and bivariate genetic correlation for a series of morphological crown variants of the anterior dentition. These results provide insight into the value of dental phenotypes as evolutionary proxies, as well as the development of tooth crowns as integrated or modular structures. Materials and methods African American dental casts from the Menegaz-Bock collection were scored for a standard set of dental morphological variables using the Arizona State Dental Anthropology System. Estimates of narrow-sense heritability and genetic correlations were generated using SOLAR v. 8.1.1, controlling for the covariates of age, sex, and birth year. Analyses were run using ordinal/continuous scale variables that were then dichotomized at various breakpoints, consistent with standard practices in dental anthropology. Results Heritability estimates were low to moderate for most traits, and lower in magnitude than those reported for odontometric data from the same study sample. Only winging, canine shoveling, and canine double shoveling returned narrow-sense heritabilities that did not differ significantly from zero. Genetic correlations were high among antimeres and metameres and low for different traits scored on the same tooth crown. These results affirm standard data cleaning practices in dental biodistance. Double shoveling was atypical in returning strong negative correlations with other traits, shoveling in particular. Conclusions Additive genetic variation contributes to dental morphological variation, although the estimates are uniformly lower than those observed for odontometrics. Patterns of genetic correlation affirm most standard practices in dental biodistance. Patterns of negative pleiotropy involving lingual and labial crown features suggest a genetic architecture and developmental complex that differentially constrain morphological variation of distinct surfaces of the same tooth crown. These patterns warrant greater consideration and cross-population validation.

Objectives This study investigates where the Xujiayao juvenile (I1 and C1) fits into the array of perikymata distribution patterns found within the genus Homo. Materials and Methods In addition to the I1 and the C1 of the Xujiayao juvenile, this study includes samples of early Homo (H. rudolfensis and H. erectus), Neandertals, early modern humans (Qafzeh), and recent modern humans from Southern Africa, Newcastle (UK), and North America (Inupiaq, AK). Three sets of analyses were undertaken, including a comparison of percentage of perikymata in the cervical half of the crown, repeated measures analysis of the percentage of total perikymata in each decile, and canonical variates analysis using both total perikymata number and the percentage of perikymata in the cervical half of the crown. Results The I1 and C1 of early Homo and Neandertals have a lower percentage of perikymata in the cervical half of the crown than modern human samples. Repeated measures analysis reveals clear distinctions in the distribution of perikymata between the modern human and fossil samples. Canonical variates analysis suggests greater differences between modern humans and the fossil samples than within the fossil samples, and classifies the Xujiayao teeth among modern humans. Discussion The present study further clarifies variation of perikymata distribution patterns within the genus Homo. The perikymata distribution of the Xujiayao juvenile tends to be more similar to that of modern humans than to either early Homo or Neandertals. Am J Phys Anthropol 157:684–693, 2015. © 2015 Wiley Periodicals, Inc.

This study investigates whether variants in dental morphology and nuclear DNA provide similar patterns of intergroup affinity among regional populations using biological distance (biodistance) estimates. Many biodistance studies of archaeological populations use skeletal variants in lieu of ancient DNA, based on the widely accepted assumption of a strong correlation between phenetic- and genetic-based affinities. Within studies of dental morphology, this assumption has been well supported by research on a global scale but remains unconfirmed at a more geographically restricted scale. Paired genetic (42 microsatellite loci) and dental (nine crown morphology traits) data were collected from 295 individuals among four contemporary Kenyan populations, two of which are known ethnically as “Swahili” and two as “Taita;” all have welldocumented population histories. The results indicate that biodistances based on genetic data are correlated with those obtained from dental morphology. Specifically, both distance matrices indicate that the closest affinities are between population samples within each ethnic group. Both also identify greater divergence among samples from the different ethnic groups. However, for this particular study the genetic data may provide finer resolution at detecting overall among-population relationships. Am J Phys Anthropol 000:000–000, 2015. VC 2015 Wiley Periodicals, Inc.

Evidence of a periodic biorhythm is retained in tooth enamel in the form of Retzius lines. The periodicity of Retzius lines (RP) correlates with body mass and the scheduling of life history events when compared between some mammalian species. The correlation has led to the development of the inter-specific Havers–Halberg oscillation (HHO) hypothesis, which holds great potential for studying aspects of a fossil species biology from teeth. Yet, little is known about the potential role of the HHO for human skeletal growth. Here, we explore this hypothesis within a sample of human skeletons. Associations are sought between the biorhythm and two hard tissues that form at different times during human ontogeny, using standard histological methods. First, we investigate relationships of RP to permanent molar enamel thickness and the underlying daily rate that ameloblasts secrete enamel during the early childhood years. Second, we develop preliminary research previously conducted on small samples of adult human bone by testing associations between RP, adult femoral length, and the rate of osteocyte proliferation. Results reveal RP is positively correlated with enamel thickness, negatively correlated with femoral length, but weakly associated with the rate of enamel secretion and osteocyte proliferation. These new data imply that a slower biorhythm predicts thicker enamel for children but shorter stature for adults. Our results develop an intra-specific HHO hypothesis suggesting a common underlying systemic biorhythm has a role in the final products of human enamel thickness and femoral length, probably through the duration rather than the rate of growth.

Here we describe dental remains from a Neanderthal fossil assemblage from Moula-Guercy, France. Our report demonstrates that the Moula-Guercy hominid remains contribute important morphological, developmental, and behavioral data to understanding Neanderthal evolutionary history. We include gross comparative morphological descriptions and enamel surface microstructure and microwear data. These teeth reveal numerous characteristics that are diagnostic of Neanderthals and provide no evidence for the presence of any other hominid taxa. Enamel growth increment data from the Moula-Guercy specimens yield evidence of a Neanderthal pattern of development, although at the lower end of the range of variation. The presence of a significant number of linear enamel hypoplasias indicates that these individuals were stressed during childhood. Molar microwear data suggest that these Neanderthals did not differ significantly from modern humans in terms of the fracture properties of the food they were consuming. The incisor microwear and macro striations provide evidence that these individuals may have been using their anterior teeth as tools, similar to the practices of several modern human populations such as the Inuit, Ipiutak, and Australian Aboriginals, and reminiscent of evidence from other Neanderthals from Krapina, Croatia, as well as the 600,000 year old hominids from Sima de los Huesos, Spain. Am J Phys Anthropol 151:477-491, 2013.© 2013 Wiley Periodicals, Inc.

The patterning cascade model of tooth morphogenesis has emerged as a useful tool in explaining how tooth shape develops and how tooth evolution may occur. Enamel knots, specialized areas of dental epithelium where cusps initiate, act as signaling centers that direct the growth of surrounding tissues. For a new cusp to form, an enamel knot must form beyond the inhibition fields of other enamel knots. The model predicts that the number and size of cusps depends on the spacing between enamel knots, reflected in the spacing between cusps. Recently, work by our group demonstrated that the model predicted Carabelli trait expression in human first molars. Here we test whether differences in Carabelli trait expression along the molar row can also be predicted by the model. Crown areas and intercusp distances were measured from dental casts of 316 individuals with a digital microscope. Although absolute cusp spacing is similar in first and second molars, the smaller size and more triangular shape of second molars results in larger cusp spacing relative to size and, likely, less opportunity for the Carabelli trait to form. The presence and size of the hypocone (HY) and a range of small accessory cusps in a larger sample of 340 individuals were also found to covary with the Carabelli trait in a complex way. The results of this study lend further support to the view that the dentition develops, varies, and evolves as a single functional complex.

Across mammalian species, the periodicity with which enamel layers form (Retzius periodicity) in permanent teeth corresponds with average body mass and the pace of life history. According to the Havers-Halberg Oscillation hypothesis (HHO), Retzius periodicity (RP) is a manifestation of a biorhythm that is also expressed in lamellar bone. Potentially, these links provide a basis for investigating aspects of a species' biology from fossilized teeth. Here, we tested intra-specific predictions of this hypothesis on skeletal samples of human juveniles. We measured daily enamel growth increments to calculate RP in deciduous molars (n = 25). Correlations were sought between RP, molar average and relative enamel thickness (AET, RET), and the average amount of primary bone growth (n = 7) in humeri of age-matched juveniles. Results show a previously undescribed relationship between RP and enamel thickness. Reduced major axis regression reveals RP is significantly and positively correlated with AET and RET, and scales isometrically. The direction of the correlation was opposite to HHO predictions as currently understood for human adults. Juveniles with higher RPs and thicker enamel had increased primary bone formation, which suggests a coordinating biorhythm. However, the direction of the correspondence was, again, opposite to predictions. Next, we compared RP from deciduous molars with new data for permanent molars, and with previously published values. The lowermost RP of 4 and 5 days in deciduous enamel extends below the lowermost RP of 6 days in permanent enamel. A lowered range of RP values in deciduous enamel implies that the underlying biorhythm might change with age. Our results develop the intra-specific HHO hypothesis.

Previous studies have suggested that Neandertals experienced greater physiological stress and/or were less capable of mitigating stress than most prehistoric modern human populations. The current study compares estimates of dental fluctuating asymmetry (DFA) for prehistoric Inupiat from Point Hope Alaska, the Late Archaic, and Protohistoric periods from Ohio and West Virginia, and a modern sample from Ohio to Neandertals from Europe and Southwest Asia. DFA results from developmental perturbation during crown formation and is thus an indicator of developmental stress, which previous studies have found to be higher in Neandertals than in several modern human populations. Here, we use recent methodological improvements in the analysis of fluctuating asymmetry suggested by Palmer and Strobeck (Annu Rev Ecol Syst 17 (1986) 391-421, Developmental instability: causes and consequences (2003a) v.1-v.36, Developmental instability: causes and consequences (2003b) 279-319) and compare the fit of Neandertal DFA Index values with those of modern humans. DFA estimates for each of the modern population samples exceeded measurement error, with the Inupiat exhibiting the highest levels of DFA for most tooth positions. All significant Neandertal z-scores were positive, exceeding the estimates for each of the modern prehistoric groups. Neandertals exhibited the fewest significant differences from the Inupiat (9.2% of values are significant at P < 0.05), while for the other modern prehistoric groups more than 10% of the Neandertal z-scores are significant at P < 0.05, more than 90% of these significant scores at P < 0.01. These results suggest that the Inupiat experienced greater developmental stress than the other prehistoric population samples, and that Neandertals were under greater developmental stress than all other prehistoric modern human samples.

Physiological stress, such as malnutrition or illness, can disrupt normal enamel growth, resulting in linear enamel hypoplasias (LEHs). Although ecological factors may contribute to LEH expression, other factors, such as surface abrasion and enamel growth variables, are also likely to be involved. Attention to these other factors is necessary before we can begin to understand what LEH might signify in terms of ecological sources of physiological stress in non-human primates. This study focuses on assessing the contribution of these other factors to variation in LEH expression within and across great ape taxa. Here, we present LEH data from unabraded crown regions in samples of seven great ape species. We analyze these data with respect to lateral enamel formation time and the angles that striae of Retzius make with the enamel surface, as these variables are expected to affect variation in LEH expression. We find that although the duration of enamel formation is associated with sex differences in LEH expression, it is not clearly related to taxonomic variation in LEH expression, and does not explain the low frequency of LEH in mountain gorillas found in this and a previous study. Our data on striae of Retzius angles suggest that these influence LEH expression along the tooth crown and may contribute to the consistently high frequencies of LEH seen in Pongo in this and previous studies. We suggest that future work aimed at understanding species variation in these angles is crucial to evaluating taxonomic patterns of LEH expression in great apes.

Previous research has demonstrated that great ape and macaque males achieve large canine crown sizes primarily through extended canine growth periods. Recent work has suggested, however, that platyrrhine males may achieve larger canine sizes by accelerating rather than prolonging growth. This study tested the hypothesis that the ontogenetic pathway leading to canine sexual dimorphism in catarrhines differs from that of platyrrhines. To test this hypothesis, males and females of several catarrhine genera (Hylobates, Papio, Macaca, Cercopithecus, and Cercocebus) and three platyrrhine genera (Cebus, Ateles, and Callicebus) were compared in the number and spacing of perikymata (enamel growth increments) on their canine crowns. In addition, perikymata periodicities (the number of days of growth perikymata represent) were determined for five genera (Hylobates, Papio, Macaca, Cebus, and Ateles) using previously published as well as original data gathered for this study. The central findings are as follows: 1) males have more perikymata than females for seven of eight genera (in five of the seven, the differences are statistically significant); 2) in general, the greater the degree of sexual dimorphism, the greater the sex difference in male and female perikymata numbers; 3) there is no evidence of a systematic sex difference in primate periodicities; and 4) there is some evidence that sex differences in enamel formation rates may make a minor contribution to canine sexual dimorphism in Papio and Cercopithecus. These findings strongly suggest that in both catarrhines and platyrrhines prolongation of male canine growth is the primary mechanism by which canine crown sexual dimorphism is achieved.

In this investigation, deciduous teeth (canines, c; first molars, m1; second molars, m2) and their permanent successors (canines, C; first premolars, P1; second premolars, P2) were used to test two related hypotheses about fluctuating asymmetry (FA). First, based on the biology of the developing dentition, it was predicted that deciduous teeth would be more developmentally stable and thus exhibit less dimensional FA than their permanent successors. Second, based on sex differences in tooth development, it was predicted that female canines would have greater developmental stability (less FA) than male canines. Bucco-lingual measurements were made on dental casts from a single Gullah population. Using a repeated-measures study design (n = 3 repeated measures), we tested these hypotheses on sample sizes ranging from 63-82 antimeric pairs. Neither hypothesis was supported by our data. In most cases, Gullah deciduous teeth did not exhibit statistically significantly less FA than their permanent successors; indeed, statistically significant differences were found for only 3 of 12 deciduous vs. permanent contrasts, and in two cases, the deciduous tooth had greater FA. Female mandibular canines exhibited statistically significantly greater FA than those of males, while there was no statistically significant sex difference in FA for the maxillary canine. FA in these Gullah samples is high when compared to Archaic and late prehistoric Ohio Valley Native Americans, consistent with historical and archaeological evidence that environmental stress was relatively higher in the Gullah population. We suggest that when environmental stress in a population is high, the impact of differences in tooth formation time spans and developmental buffering upon FA may be minor relative to the effect of developmental noise.

This study investigates changes in the prevalence of linear enamel hypoplasia (LEH) before and after the shift from irregular to regular provisioning in the Cayo Santiago rhesus monkey population. Prior to 1956, monkeys on this island colony did not receive consistent provisions, and were reported to be in poor health (Rawlins and Kessler [1986] The Cayo Santiago Macaques; Albany: State University of New York Press). A regular provisioning program, instituted in August 1956, resulted in the improved health of individuals and the growth of the population (Rawlins and Kessler [1986] The Cayo Santiago Macaques; Albany: State University of New York Press). LEH, a developmental defect of enamel, is a sensitive indicator of systemic physiological stress (Goodman and Rose [1990] Yrbk. Phys. Anthropol. 33:59-110). It was therefore hypothesized that the prevalence of LEH would be higher in monkeys who were irregularly provisioned than in monkeys who experienced regular provisioning. To test this hypothesis, teeth were examined for LEH in a sample of 181 female rhesus monkeys. The results support the hypothesis: the mean number of defects was statistically significantly higher in the preprovisioned group than it was in the postprovisioned one. When LEH prevalence was assessed using only defects occurring on antimeric pairs, the preprovisioned group again had a higher prevalence than the postprovisioned one, although the difference was not statistically significant, most likely because of the reduced sample size. The results of this study indicate that changes in LEH prevalence, at least in this population of rhesus monkeys, are associated with changes in nutritional status.

This study uses macroscopic and micro- scopic methods to analyze the expression of linear enamel hypoplasia (LEH) in Plio-Pleistocene South African homi- nins. LEH is a developmental defect of enamel that is used in many anthropological contexts as a physiological stress in- dicator. Previous research has not settled the question as to whether differences in LEH expression exist between Paran- thropus and Australopithecus and if they exist, to what ex- tent these differences might be explained simply by taxo- nomic differences in enamel development and morphology rather than by differential stress experience. In this study, the analysis of LEH is conducted with respect to differences between Paranthropus and Australopithecus in aspects of enamel development and morphology that are thought to influence LEH expression. Two factors impacting LEH ex- pression are considered: the duration of enamel formation, and the spacing of perikymata. It is predicted that if the first factor strongly influences the expression of LEH, then there should be fewer defects per tooth in Paranthropus because of its abbreviated crown formation spans (and fast extension rates) relative to Australopithecus. It is also predicted that because Australopithecus has more densely packed periky- mata in comparable regions of the crown than Paranthropus, this taxon should, on average, have narrower defects than Paranthropus. To address these questions, 200 Australo- pithecus and 137 Paranthropus teeth were examined for LEH, and the analysis of defect width with respect to periky- mata spacing was conducted on tooth impressions examined under a scanning electron microscope using INCA (Oxford Instruments) measurement software. Data support the first prediction: Australopithecus does have significantly more defects per canine tooth than Paranthropus. Data do not support the second prediction in large part because several Australopithecus specimens have wide groove defects in which perikymata are not visible and enamel is irregular. Such wide grooves are not predicted by perikymata spacing such that alternative explanations, including taxonomic dif- ferences in ameloblast sensitivity and the duration/severity of disruptions to enamel growth, must be considered. Am J Phys Anthropol 120:309-322, 2003. © 2003 Wiley-Liss, Inc.

This study describes the expression of linear enamel hypoplasia (LEH), a sensitive dental indicator of physiological stress, in Thailand gibbons (Hylobates lar carpenteri). Previous studies of enamel hypoplasia in hominoids have focused on great apes, with little attention given to the expression of this stress indicator in gibbons. In that gibbons differ from both monkeys and great apes in numerous life history features, LEH expression in gibbons might be expected to show significant differences from both. In this study, 92 gibbon specimens from two sites in Thailand were compared with several samples of monkeys and great apes in their expression of LEH. The intertooth distribution of LEH in gibbons was compared to that of chimpanzees and rhesus monkeys. Gibbon populations from both sites exhibit LEH frequencies intermediate between those of the monkey samples, in which LEH prevalence is usually low, and those of the great ape samples, in which LEH prevalence is high. Gibbons differ significantly from monkeys, but not great apes, in the number of individuals whose teeth record multiple stress events. Multiple episodes of stress are rarely recorded in the teeth of monkeys, while multiple stress events occur with higher frequency in gibbons and great apes. Taxonomic variation in the duration of crown formation, the prominence and spacing of perikymata on dental crowns, life history features, and/or experience of physiological stress may explain these patterns. The intertooth distribution of LEH in gibbons is, for different reasons, unlike that of either chimpanzees or rhesus monkeys. The mandibular canines of gibbons have significantly more LEH than any of their other teeth. Aspects of crown morphology, perikymata prominence/spacing, enamel thickness, and crown formation spans are potential causes of taxonomic variation in the intertooth distribution of LEH.

Three hundred and sixty rhesus macaque specimens at the Caribbean Primate Research Center were examined for evidence of linear enamel hypoplasia (LEH). A previously unreported intertooth pattern in LEH was observed. Defects occur preferentially on the sectorial premolar of both males and females. Relative to other teeth, the sectorial premolar exhibits more prominent defects and is more likely to exhibit multiple defects. This pattern is unlike the human intertooth LEH pattern and unlike patterns previously reported for monkeys and apes. These observations are discussed in the context of factors thought to influence the intertooth distribution of LEH in humans and in nonhuman primates. The authors reject crown height, the timing of crown development, and the duration of crown formation as factors contributing to the observed pattern and favor an explanation involving enamel thickness, perikymata spacing, and/or prism orientation. Am J Phys Anthropol 107:179–186, 1998. © 1998 Wiley-Liss, Inc.

This study reconstructs linear enamel hypoplasia (LEH) prevalence and stress episode duration among Jomon period foragers from Hokkaido, Japan (HKJ). Results are compared to Jomon period samples from coastal Honshu, Japan (HSJ) and Tigara Inupiat from Point Hope, Alaska (PHT) to provide a more comprehensive perspective on the manifestation of stress among circum-Pacific foragers. LEH were identified macro- and microscopically by enamel surface depressions and increased perikymata spacing within defects. Individuals with more than one anterior tooth affected by LEH were labeled as LEH positive. Stress episode durations were estimated by counting the number of perikymata within the occlusal wall of each LEH and multiplying that number by constants reflecting modal periodicities for modern human teeth. LEH prevalence and stress episode duration did not differ significantly between the two Jomon samples. Significantly greater frequencies of LEH were found in HKJ as compared to PHT foragers. However, HKJ foragers had significantly shorter stress episode durations as compared to PHT. This suggests that a greater proportion of HKJ individuals experienced stress episodes than did PHT individuals, but these stress events ended sooner. Similarity in stress experiences between the two Jomon samples and differences between the HKJ and PHT are found. These findings are important for two reasons. First, stress experiences of foraging populations differ markedly and cannot be generalized by subsistence strategy alone. Second, due to significant differences in episode duration, stress experiences cannot be understood using prevalence comparisons alone.

We describe and analyze a Neandertal postcranial skeleton and dentition, which together show unambiguous signs of right-handedness. Asymmetries between the left and right upper arm in Regourdou 1 were identified nearly 20 years ago, then confirmed by more detailed analyses of the inner bone structure for the clavicle, humerus, radius and ulna. The total pattern of all bones in the shoulder and arm reveals that Regourdou 1 was a right-hander. Confirmatory evidence comes from the mandibular incisors, which display a distinct pattern of right oblique scratches, typical of right-handed manipulations performed at the front of the mouth. Regourdou's right handedness is consistent with the strong pattern of manual lateralization in Neandertals and further confirms a modern pattern of left brain dominance, presumably signally linguistic competence. These observations along with cultural, genetic and morphological evidence indicate language competence in Neandertals and their European precursors.

Recent studies have suggested that Neandertals and modern humans differ in the distribution of perikymata (enamel growth increments) over their permanent anterior tooth crowns. In modern humans, perikymata become increasingly more compact toward the cervix than they do in Neandertals. Previous studies have suggested that a more homogeneous distribution of perikymata, like that of Neandertals, characterizes the anterior teeth of Homo heidelbergensis and Homo erectus as well. Here, we investigated whether Qafzeh anterior teeth (N = 14) differ from those of modern southern Africans, northern Europeans, and Alaskans (N = 47-74 depending on tooth type) in the percentage of perikymata present in their cervical halves. Using the normally distributed modern human values for each tooth type, we calculated Z-scores for the 14 Qafzeh teeth. All but two of the 14 Qafzeh teeth had negative Z-scores, meaning that values equal to these would be found in the bottom 50% of the modern human samples. Seven of the 14 would be found in the lowest 5% of the modern human distribution. Qafzeh teeth therefore appear to differ from those of modern humans in the same direction that Neandertals do: with generally lower percentages of perikymata in their cervical regions. The similarity between them appears to represent the retention of a perikymata distribution pattern present in earlier members of the genus Homo, but not generally characteristic of modern humans from diverse regions of the world.

Linear enamel hypoplasia (LEH), a type of enamel defect reflecting nonspecific physiological stress, has traditionally been used by bioarchaeologists to assess human health. Initially, measurements of defect width were used to estimate the duration of stress episodes. More recently, methods of counting within-defect perikymata (enamel growth increments) were developed to more accurately assess duration. Because perikymata are often not continuously visible within defects, while widths can usually be measured, the primary purpose of this article was to determine if, under restrictive conditions, the widths of LEH defects might be used as relative indicators of stress episode duration. Using a set of dental replicas from the prehistoric Irene Mound (1150-1400 A.D.), this study also investigated potential sources of variation in defect widths and how often defect widths could be measured and within-defect perikymata counted. Of 120 defects, only 47 contained both measurable defect widths and total within-defect perikymata, while 79 had measurable defect widths. Regression analysis revealed that, for these 47 defects, defect widths were more strongly related to the total number of within-defect perikymata than they were to crown region or tooth type. Although wide prediction intervals indicated that a defect's width could not be used to predict the number of within-defect perikymata for an individual, narrower confidence intervals associated with hypothetical mean population widths suggested that mean defect widths might be used to rank populations in terms of relative average stress episode duration.

Previous researchers hypothesized that tooth types forming during early childhood should be less sexually dimorphic than those forming during later childhood, if sex hormone concentration differences between males and females increase progressively throughout childhood and can affect tooth size. Descriptive tooth size data have recently been cited in support of this hypothesis, particularly with respect to differences in sexual dimorphism among the tooth types of tooth classes. The present study tests this hypothesis for the mesiodistal dimension of human permanent teeth using published data for incisor, premolar, and molar tooth classes from seven diverse populations. The sample size for each tooth type per population was at least 50. This study also tests a modification of this hypothesis which takes into account the postnatal testosterone surge in males and the low levels of sex hormones in both sexes prior to puberty. Predictions are developed for both the original and modified hypotheses. The "D" statistic, the total area of nonoverlap between the phenotypic distributions of males and females, is used to quantify sexual dimorphism. Comparison of D values for different tooth types within tooth classes across these seven populations does not strongly support either hypothesis. These results suggest that gross changes in sex hormone concentrations during development are not related to population-wide patterns of sexual dimorphism among the tooth types of human permanent tooth classes, as recent studies indicate. This finding is consistent with other studies which suggest that sex hormones have only a minor role in generating crown size sexual dimorphism.

Linear enamel hypoplasia (LEH) is an enamel defect that records the effects of physiological stress on tooth formation. Estimating the age at which LEH defects form is integral to the reconstruction of population health in bioarcheological studies. Two principal methods for aging LEH defects have been introduced in the literature. The conventional approach employs regression equations based on a linear model of tooth growth. The newer, Reid and Dean [Am J Phys Anthropol 113 (2000) 135-139] approach, is based upon a histologically derived curvilinear model of enamel development and therefore likely provides more accurate age estimates. However, the extent to which the Reid and Dean method produces estimated ages at defect formation differing from those of the regression equations has not, until now, been determined. This study quantifies the differences between these two methods. Evaluating the degree to which these methods differ is essential for interpreting the accuracy of LEH age estimates given in previous bioarcheological studies. Age estimates of LEH defects on 338 anterior teeth from the Hamann-Todd osteological sample were calculated using both methods. The resulting estimated ages were compared through a randomized block ANOVA. However, the mean differences between the estimated ages yielded by both methods range from 4 months or less depending on the tooth type with an overall average of 2.63 months. The discussion focuses on the degree to which this difference affects answers to bioarcheological questions.

Linear enamel hypoplasia (LEH), a developmental defect of enamel, increases in frequency from prosimian to monkey to lesser ape to great ape grades (Guatelli-Steinberg 2000 Am. J. Phys. Anthropol. 112:395-410, [2001] Evol. Anthropol. 10:138-151; Newell 1998 Ph.D. dissertation, Temple University). This taxonomic pattern in the distribution of LEH is closely related to maturation length across the primate order (Newell 1998 Ph.D. dissertation, Temple University, 2000 Am. J. Phys. Anthropol. [Suppl.] 30:236). Longer maturation periods are associated with higher LEH frequencies; they appear to provide greater opportunity for defects to form. The present study explores the relationship between maturation length and LEH frequency within the Ceboidea. Because of its prolonged period of growth, Cebus is predicted to manifest LEH at a higher frequency than the more rapidly maturing ceboid genera. To test this hypothesis, two separate researchers (E.A.N. and D.G.-S.) examined LEH in nonoverlapping museum series of ceboids. The results support the hypothesis: in 13 genera (n = 1,276), E.A.N. found that LEH frequencies ranged from 0% in Callicebus, Cebuella, and Saimiri to 20% in Cebus. D.G.-S. found similar frequencies among five genera (n = 107), from 0% in Saimiri to 32% in Cebus. Thus, the broad pattern of LEH distribution evident across major taxonomic groups of primates is repeated within the Ceboidea. We also examined a related hypothesis linking the spacing of perikymata, which is influenced by enamel extension rates (Shellis 1998 J. Hum. Evol. 35:387-400), to LEH. The most likely areas of tooth crowns to exhibit LEH in human teeth are those in which perikymata are most closely spaced (Hillson and Bond 1997 Am. J. Phys. Anthropol. 104:89-103). We hypothesized that the longer-maturing Cebus, with its elevated LEH frequency, will also exhibit more closely spaced perikymata than other ceboids. Analysis of a small microscopic subsample (n = 8) lends limited support to this second hypothesis.

This report documents greater variabil- ity in early hominin first molar dental trait frequencies than previous research indicated. Specifically, frequen- cies of several M1 dental traits that previously appeared to uniquely characterize Paranthropus are shown here to resemble those of the A. afarensis sample from Hadar. Like Paranthropus, A. afarensis from Hadar has a high frequency of cusp 6 and a low frequency of the proto- stylid. Paranthropus and A. afarensis are also not statis- tically significantly different in their frequencies of LM1 cusp 7, although this cusp is actually absent in the lat- ter. Both groups differ significantly from A. africanus in their frequencies of these traits. Based on the develop- mental biology of molar cusp patterns, we suggest that the morphological similarities between Paranthropus and the Hadar sample may be homoplasies. Am J Phys Anthropol 127:000-000, 2005. V

Most studies report a high prevalence of linear enamel hypoplasia (LEH) in the great apes relative to other nonhuman primates and some human populations. It is unclear if this difference is a direct result of poor health status for the great apes, or if it represents differential incidence due to a lower threshold (sensu Goodman and Rose, 1990 Am. J. Phys. Anthropol. [suppl.] 33:59-110) for the occurrence of enamel hypoplasia among great apes. This study uses the Smithsonian National Museum of Natural History's great ape collection to examine the prevalence of LEH, the most common type of hypoplasia observed. Frequencies of LEH are reported, as well as analyses by taxa and provenience. The study sample consists of 136 specimens and includes 41 gorillas, 25 chimpanzees, and 70 orangutans. Analyses of frequencies are presented for both individuals and teeth by taxonomic category and locality. Among the individuals in this study, 63.97% are affected by LEH. Overall, gorillas (29.27%) exhibit lower frequencies of LEH than chimpanzees (68.00%) and orangutans (82.86%). There is a marked difference in LEH frequencies between mountain and lowland gorillas. There is no difference in LEH frequencies between Sumatran and Bornean orangutans. A range of variation for the great apes in enamel hypoplasia frequencies is found when taxon and locality are considered. It is likely that both biological and environmental factors influence the high frequencies of enamel hypoplasia exhibited in the great apes.

This study of linear enamel hypoplasia (LEH) in Plio-Pleistocene hominins builds on a previous study (Guatelli-Steinberg [2003] Am. J. Phys. Anthropol. 120:309-322) that focused on LEH in early South African hominins. The present study is more comprehensive, encompassing dental specimens of hominins from East Africa as well, including early Homo. As a developmental defect of enamel, LEH is used in anthropological contexts to reveal information about physiological stress. However, intrinsic aspects of enamel development and morphology can affect the expression of LEH, complicating efforts to understand the significance of these defects. In this study, the analysis of LEH is conducted with respect to enamel development and morphology. It is predicted that Paranthropus should have fewer defects on its canine teeth than Australopithecus and Homo, owing to its abbreviated period of enamel formation. This prediction is supported: Paranthropus has statistically significantly fewer defects per canine than Australopithecus and Homo. The previous study demonstrated that despite the wider spacing of perikymata on the teeth of South African Paranthropus, defects on the canine teeth of this genus were not wider than those of Australopithecus. A multiple linear regression analysis in that study, as well as a separate analysis in the present study, indicate that the number of perikymata within defects is a better predictor of defect width than perikymata spacing. In this study, it was additionally found that the average number of perikymata within Australopithecus defects is statistically significantly greater than it is in Paranthropus, thus explaining why Paranthropus defects are not wider than those of Australopithecus. The biological significance of this difference in the number of perikymata within the defects of Australopithecus and Paranthropus is considered in light of several factors, including: 1) the possibility that other intrinsic attributes of enamel morphology may be involved (specifically the faster extension rates of Paranthropus that result in shallower defects), 2) generic differences in the canalization of enamel development, and 3) generic differences in the duration of disruptions to enamel growth.

Background The patterning cascade model of tooth morphogenesis accounts for shape development through the interaction of a small number of genes. In the model, gene expression both directs development and is controlled by the shape of developing teeth. Enamel knots (zones of nonproliferating epithelium) mark the future sites of cusps. In order to form, a new enamel knot must escape the inhibitory fields surrounding other enamel knots before crown components become spatially fixed as morphogenesis ceases. Because cusp location on a fully formed tooth reflects enamel knot placement and tooth size is limited by the cessation of morphogenesis, the model predicts that cusp expression varies with intercusp spacing relative to tooth size. Although previous studies in humans have supported the model's implications, here we directly test the model's predictions for the expression, size, and symmetry of Carabelli cusp, a variation present in many human populations. Methodology/Principal Findings In a dental cast sample of upper first molars (M1s) (187 rights, 189 lefts, and 185 antimeric pairs), we measured tooth area and intercusp distances with a Hirox digital microscope. We assessed Carabelli expression quantitatively as an area in a subsample and qualitatively using two typological schemes in the full sample. As predicted, low relative intercusp distance is associated with Carabelli expression in both right and left samples using either qualitative or quantitative measures. Furthermore, asymmetry in Carabelli area is associated with asymmetry in relative intercusp spacing. Conclusions/Significance These findings support the model's predictions for Carabelli cusp expression both across and within individuals. By comparing right-left pairs of the same individual, our data show that small variations in developmental timing or spacing of enamel knots can influence cusp pattern independently of genotype. Our findings suggest that during evolution new cusps may first appear as a result of small changes in the spacing of enamel knots relative to crown size.