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New methodology to reconstruct in 2-D the cuspal enamel of modern human lower molars
Abstract and Figures
Objectives:
In the last years different methodologies have been developed to reconstruct worn teeth. In this article, we propose a new 2-D methodology to reconstruct the worn enamel of lower molars. Our main goals are to reconstruct molars with a high level of accuracy when measuring relevant histological variables and to validate the methodology calculating the errors associated with the measurements.
Methods:
This methodology is based on polynomial regression equations, and has been validated using two different dental variables: cuspal enamel thickness and crown height of the protoconid. In order to perform the validation process, simulated worn modern human molars were employed. The associated errors of the measurements were also estimated applying methodologies previously proposed by other authors.
Results:
The mean percentage error estimated in reconstructed molars for these two variables in comparison with their own real values is -2.17% for the cuspal enamel thickness of the protoconid and -3.18% for the crown height of the protoconid. This error significantly improves the results of other methodologies, both in the interobserver error and in the accuracy of the measurements.
Conclusions:
The new methodology based on polynomial regressions can be confidently applied to the reconstruction of cuspal enamel of lower molars, as it improves the accuracy of the measurements and reduces the interobserver error. The present study shows that it is important to validate all methodologies in order to know the associated errors. This new methodology can be easily exportable to other modern human populations, the human fossil record and forensic sciences.
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... presence and extent of wear (Guatelli-Steinberg, 2008;Guatelli-Steinberg, Ferrell, & Spence, 2012;Lockey, Alemseged, Hublin, & Skinner, 2020;Modesto-Mata et al., 2017O'Hara, Le Cabec, Xing, Skinner, & Guatelli-Steinberg, 2019;Saunders, Chan, Kahlon, Kluge, & FitzGerald, 2007;Skinner, Alemseged, Gaunitz, & Hublin, 2015;Smith, Kupczik, Machanda, Skinner, & Zermeno, 2012). To measure critical dental growth and enamel thickness variables, dental researchers require unworn teeth and/or accurate and reliable methods to reconstruct worn tooth crowns (e.g., Guatelli-Steinberg, 2008;Guatelli-Steinberg, Pampush, et al., 2018;Lockey et al., 2020;Martin, 1985;Modesto-Mata et al., 2017;Saunders et al., 2007;Schwartz, 2000;Skinner et al., 2015). ...
... presence and extent of wear (Guatelli-Steinberg, 2008;Guatelli-Steinberg, Ferrell, & Spence, 2012;Lockey, Alemseged, Hublin, & Skinner, 2020;Modesto-Mata et al., 2017O'Hara, Le Cabec, Xing, Skinner, & Guatelli-Steinberg, 2019;Saunders, Chan, Kahlon, Kluge, & FitzGerald, 2007;Skinner, Alemseged, Gaunitz, & Hublin, 2015;Smith, Kupczik, Machanda, Skinner, & Zermeno, 2012). To measure critical dental growth and enamel thickness variables, dental researchers require unworn teeth and/or accurate and reliable methods to reconstruct worn tooth crowns (e.g., Guatelli-Steinberg, 2008;Guatelli-Steinberg, Pampush, et al., 2018;Lockey et al., 2020;Martin, 1985;Modesto-Mata et al., 2017;Saunders et al., 2007;Schwartz, 2000;Skinner et al., 2015). ...
... Three methods for reconstructing cusp tips have been established by different groups of researchers and on different taxa. In this study, the three methods are referred to as the "Pen Tool" method (Guatelli-Steinberg, Ferrell, & Spence, 2012;O'Hara et al., 2019;Saunders et al., 2007), the "Polynomial" method (Modesto-Mata et al., 2017), and the "Profile" method (Grine & Martin, 1988;Martin, 1983;Smith et al., 2011;Smith, Kupczik, et al., 2012). The present article offers the first comparative study of the three methods on a single sample, allowing for a direct comparison of their accuracy across four measurements: crown height (CH), cuspal enamel thickness (CET), regional enamel thickness (RegAET), and average enamel thickness (AET). ...
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.
... Worn crowns were reconstructed following the recommendations of O'Hara and Guatelli-Steinberg (2021). These authors found that for AET and crown height measurements, crowns with limited wear (for which wear did not reach the dentine horns and/or the deepest point of the occlusal basin) accurate values were achievable with the Profile(Grine & Martin, 1988;Smith et al., 2011;Smith et al., 2012), Polynomial(Modesto-Mata et al., 2017), and Pen Tool methods(Guatelli-Steinberg et al., 2009;O'Hara et al., 2019;Saunders et al., 2007). O'Hara and Guatelli-Steinberg (2021) also found that, for crowns on which wear exposed the tips of dentine horns (what they termed "extensive wear"), it was still possible to obtain accurate AET values using the Profile or Polynomial methods. ...
To test the hypothesis that differences in crown structure, enamel growth, and crown geometry in Cercocebus and Lophocebus molars covary with differences in the feeding strategies (habitual vs. fallback durophagy, respectively) of these two genera. Relative to Lophocebus molars, Cercocebus molars are predicted to possess features associated with greater fracture resistance and to differ in enamel growth parameters related to these features. Sample proveniences are as follows: Cercocebus atys molars are from the Taï Forest, Ivory Coast; Lophocebus albigena molars are from a site north of Makoua, Republic of Congo; and a Lophocebus atterimus molar is from the Lomako Forest, Democratic Republic of Congo. For μCT scans on which aspects of molar form were measured, sample sizes ranged from 5 to 35 for Cercocebus and 3 to 12 for Lophocebus. A subsample of upper molars was physically sectioned to measure enamel growth variables. Partly as a function of their larger size, Cercocebus molars had significantly greater absolute crown strength (ACS) than Lophocebus molars, supporting the hypothesis. Greater crown heights in Cercocebus are achieved through faster enamel extension rates. Also supporting the hypothesis, molar flare and proportional occlusal basin enamel thickness were significantly greater in Cercocebus. Relative enamel thickness (RET), however, was significantly greater in Lophocebus. If ACS is a better predictor of fracture resistance than RET, then Cercocebus molars may be more fracture resistant than those of Lophocebus. Greater molar flare and proportional occlusal basin thickness might also afford Cercocebus molars greater fracture resistance.
... This method has been demonstrated to reliably reconstruct cusp tips with different degrees of wear (<2% error; Saunders et al., 2007). Some studies suggest that this method is more subjective and produces higher error compared to more recently-developed polynomial regression-based methods to reconstruct worn cusps (Modesto-Mata et al., 2017), but for our purposes of estimating EDJ length and locating crown regions along the EDJ, the former method suffices. ...
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.
... For the sample of H. naledi, a more precise method was used following that of Saunders et al. (2007) and originally designed for use in histological canine sections. (A recent study has also focused on reconstructing worn crowns, though in human molars; Modesto-Mata et al., 2017.) In the Saunders et al. (2007) method, tangent lines are drawn with the line tool in Adobe Photoshop ® on either side of the crown. ...
Perikymata, incremental growth lines visible on tooth enamel surfaces, differ in their distribution and number among hominin species, although with overlapping patterns. This study asks: (1) How does the distribution of perikymata along the lateral enamel surface of Homo naledi anterior teeth compare to that of other hominins? (2) When both perikymata distribution and number are analyzed together, how distinct is H. naledi from other hominins? A total of 19 permanent anterior teeth (incisors and canines) of H. naledi were compared, by tooth type, to permanent anterior teeth of other hominins: Australopithecus afarensis, Australopithecus africanus, Paranthropus robustus, Paranthropus boisei, Homo ergaster/Homo erectus, other early Homo, Neandertals, and modern humans, with varying sample sizes. Repeated measures analyses of the percentage of perikymata per decile of reconstructed crown height yielded several statistically significant differences between H. naledi and other hominins. Canonical variates analysis of percentage of perikymata in the cervical half of the crown together with perikymata number revealed that, in 8 of 19 cases, H. naledi teeth were significantly unlikely to be classified as other hominins, while exhibiting least difference from modern humans (especially southern Africans). In a cross-validated analysis, 68% of the H. naledi teeth were classified as such, while 32% were classified as modern human (most often southern African). Of 313 comparative teeth use for this analysis, only 1.9% were classified as H. naledi. What tends to differentiate H. naledi anterior tooth crowns from those of most other hominins, including some modern humans, is strongly skewed perikymata distributions combined with perikymata numbers that fall in the middle to lower ranges of hominin values. H. naledi therefore tends toward a particular combination of these features that is less often seen in other hominins. Implications of these data for the growth and development of H. naledi anterior teeth are considered.
We present a novel method to estimate original crown height (OCH) for worn human mandibular canines using a cubic regression equation based on ratios of worn crown height and exposed dentin. This method may help alleviate issues frequently presented by worn teeth in dental analyses, including those in bioarchaeology. Mandibular canines (n = 28) from modern day New Zealand and English populations were selected. Crown height and dentin thickness were measured on dental thin sections (n = 19) and the resulting (log10) ratios were fitted to a cubic regression curve allowing OCH in worn crowns to be predicted. Variation in the dentin apex position was recorded and effects of angled wear slopes investigated allowing adjusted values to be generated. Our method is trialed for use on intact and sectioned teeth (n = 17). A cubic regression curve best describes the relationship between (log10) ratios and crown height deciles (R² = 0.996, df1 = 3, df2 = 336, p < 0.001). No significant differences were detected between OCH estimates using our method and digitally recreated cusp outlines of the same crowns (t = 1.024, df = 16, p > 0.05), with a mean absolute error of 0.171 mm and an adjusted coefficient of determination of 0.923. Our approach offers a quantitative method to estimate the percentage of OCH remaining on worn mandibular canines, and by extension, the OCH. Our estimates are comparable to digitally recreated cusps but less subjective and not limited to crowns with minimal wear.
Characterizing dental development in fossil hominins is important for distinguishing between them and for establishing where and when the slow overall growth and development of modern humans appeared. Dental development of australopiths and early Homo was faster than modern humans. The Atapuerca fossils (Spain) fill a barely known gap in human evolution, spanning ~1.2 to ~0.4 million years (Ma), during which H. sapiens and Neandertal dental growth characteristics may have developed. We report here perikymata counts, perikymata distributions and periodicities of all teeth belonging to the TE9 level of Sima del Elefante, level TD6.2 of Gran Dolina (H. antecessor) and Sima de los Huesos. We found some components of dental growth in the Atapuerca fossils resembled more recent H. sapiens. Mosaic evolution of perikymata counts and distribution generate three distinct clusters: H. antecessor, Sima de los Huesos and H. sapiens.
Previous mitochondrial DNA analyses on ancient European remains have suggested that the current distribution of haplogroup H was modeled by the expansion of the Bell Beaker culture (ca 4,500-4,050 years BP) out of Iberia during the Chalcolithic period. However, little is known on the genetic composition of contemporaneous Iberian populations that do not carry the archaeological tool kit defining this culture. Here we have retrieved mitochondrial DNA (mtDNA) sequences from 19 individuals from a Chalcolithic sample from El Mirador cave in Spain, dated to 4,760-4,200 years BP and we have analyzed the haplogroup composition in the context of modern and ancient populations. Regarding extant African, Asian and European populations, El Mirador shows affinities with Near Eastern groups. In different analyses with other ancient samples, El Mirador clusters with Middle and Late Neolithic populations from Germany, belonging to the Rössen, the Salzmünde and the Baalberge archaeological cultures but not with contemporaneous Bell Beakers. Our analyses support the existence of a common genetic signal between Western and Central Europe during the Middle and Late Neolithic and points to a heterogeneous genetic landscape among Chalcolithic groups.
The application of microtomography (mCT) to dental morphological studies has unveiled a new source of palaeobiological information, particularly in the analysis of the internal structures of teeth. In this study, we assess the expression of talonid crests at the enamel and dentine surfaces in lower permanent and second deciduous molars (M2 and dm2) of H. sapiens, H. neanderthalensis and Atapuerca-Sima de los Huesos (SH) hominins. In modern humans, talonid crests are described exclusively in the deciduous teeth (Korenhof, 1982) and interpreted as a primitive mammalian remnant of the talonid attachment to the trigonid. Here we report for the first time the expression of talonid crests of deciduous and permanent molars in H. sapiens, H. neanderthalensis and Middle Pleistocene hominins. We discuss possible evolutionary interpretations and suggest the importance of recording this feature in future studies.
Trigonid crest patterning in lower molars is distinctive among Late Pleistocene hominins such as Homo neanderthalensis, fossil Homo sapiens and modern humans. In this paper, we present an examination of trigonid crest patterning in the Middle Pleistocene permanent lower molar sample (n=62) of Homo heidelbergensis from Sima de los Huesos (SH). Crest expression was assessed from 3D models of the enamel and the dentine surfaces that were produced using micro-computed tomography (microCT). The aims of our analysis are to: 1) characterize the pattern of trigonid crest expression at the outer enamel and enamel-dentine junction surfaces (OES and EDJ) of the SH sample, 2) evaluate the concordance of expression between both surfaces, and 3) place trigonid crest variation in the SH sample into a phylogenetic context. Our results reveal a greater variability in the expression of trigonid crests at the EDJ (14 types) compared to the OES (4 types). Despite this variability, in almost all cases the expression of a continuous mid-trigonid or distal crest at the OES corresponds with the expression of a continuous mesial/mid-trigonid or distal trigonid crest, respectively, at the EDJ. Thus, it is possible to predict the type of trigonid crest pattern that would be at the OES in the case of partially worn teeth. Our study points to increased variability in trigonid crest expression in M3s compared to M1s and M2s. Moreover, our analysis reveals that the SH sample matches broadly the trigonid crest patterns displayed by H.neanderthalensis and differs from those exhibited by H.sapiens, particularly in the almost constant expression of a continuous middle trigonid crest at the EDJ. However, SH hominins also exhibit patterns that have not been reported in H.neanderthalensis and H.sapiens samples. Other aspects of the variability of the trigonid crest expression at the dentine are presented and discussed.
A modern human-like sequence of dental development, as a proxy for the pace of life history, is regarded as one of the diagnostic hallmarks of our own genus Homo1-3. Brain size, age at first reproduction, lifespan and other life-history traits correlate tightly with dental development4-6. Here we report differences in enamel growth that show the earliest fossils attributed to Homo do not resemble modern humans in their development. We used daily incremental markings in enamel to calculate rates of enamel formation in 13 fossil hominins and identified differences in this key determinant of tooth formation time. Neither australopiths nor fossils currently attributed to early Homo shared the slow trajectory of enamel growth typical of modern humans; rather, both resembled modern and fossil African apes. We then reconstructed tooth formation times in australopiths, in the ∼1.5-Myrold Homo erectus skeleton from Nariokotome, Kenya7, and in another Homo erectus specimen, Sangiran S7-37 from Java8. These times were shorter than those in modern humans. It therefore seems likely that truly modern dental development emerged relatively late in human evolution.
Two molars recovered at Trinil, Java, have been the subject of more than a century of debate since their discovery by Eugène Dubois in 1891–92. These molars have been a;ributed to several ape and human taxa (including Pan and Meganthropus), although most studies agree that they are either fossil Pongo or Homo erectus molars. Compli‑ cating the assessment of these molars is the metric and morphological similarity of Pongo and Homo erectus mo‑ lars, and uncertainty regarding their serial positions within the maxillary row. Here we applied non‑destructive conventional and synchrotron microtomographic imaging to measure the structure of these molars and aspects of their development. Comparisons were made with modern Homo and Pongo maxillary molars, as well as small samples of fossil Pongo and Homo erectus molars. Root spread was calculated from three‑dimensional surface mod‑ els, and enamel thickness and enamel‑dentine junction morphology were assessed from virtual planes of section. Developmental features were investigated using phase contrast X‑ray synchrotron imaging. PaleoAnthropology 2009: 117−129.
One highly debated issue in palaeoanthropology is that of modern human origins, particularly the issue of when 'anatomically modern humans' (AMH) from the African Middle Stone Age became fully modern. While studies of cranial and external dental morphology suggest a modern transition occurred 150 000-200 000 years ago, little is known about dental development or enamel thickness in AMH. Studies of early members of the genus Homo suggest that the modern, prolonged condition of tooth growth arose late in human evolution, and that the enamel thickness of earlier hominins may not be homologous to the modern condition. This study represents the first integrated investigation of molar crown enamel thickness, volume, and development in fossil hominins, aimed at determining whether differences between AMH and living populations can be detected in these traits. Using high-resolution micro-computed tomography, we demonstrate similarities in enamel thickness and crown volumes between fossil and modern populations. Additionally, long-period growth line numbers and estimates of crown formation times for AMH molars fall within modern human ranges. These findings suggest that tooth structure and growth have remained constant for more than 60 000 years, despite the known geographical, technological, and ecological diversity that characterizes later stages of human evolution.
Thick molar enamel is among the few diagnostic characters of hominins which are measurable in fossil specimens. Despite a long history of study and characterization of Paranthropus molars as relatively 'hyper-thick', only a few tooth fragments and controlled planes of section (designed to be proxies of whole-crown thickness) have been measured. Here, we measure molar enamel thickness in Australopithecus africanus and Paranthropus robustus using accurate microtomographic methods, recording the whole-crown distribution of enamel. Both taxa have relatively thick enamel, but are thinner than previously characterized based on two-dimensional measurements. Three-dimensional measurements show that P. robustus enamel is not hyper-thick, and A. africanus enamel is relatively thinner than that of recent humans. Interspecific differences in the whole-crown distribution of enamel thickness influence cross-sectional measurements such that enamel thickness is exaggerated in two-dimensional sections of A. africanus and P. robustus molars. As such, two-dimensional enamel thickness measurements in australopiths are not reliable proxies for the three-dimensional data they are meant to represent. The three-dimensional distribution of enamel thickness shows different patterns among species, and is more useful for the interpretation of functional adaptations than single summary measures of enamel thickness.
Humans have an unusual life history, with an early weaning age, long childhood, late first reproduction, short interbirth intervals, and long lifespan. In contrast, great apes wean later, reproduce earlier, and have longer intervals between births. Despite 80 y of speculation, the origins of these developmental patterns in Homo sapiens remain unknown. Because they record daily growth during formation, teeth provide important insights, revealing that australopithecines and early Homo had more rapid ontogenies than recent humans. Dental development in later Homo species has been intensely debated, most notably the issue of whether Neanderthals and H. sapiens differ. Here we apply synchrotron virtual histology to a geographically and temporally diverse sample of Middle Paleolithic juveniles, including Neanderthals, to assess tooth formation and calculate age at death from dental microstructure. We find that most Neanderthal tooth crowns grew more rapidly than modern human teeth, resulting in significantly faster dental maturation. In contrast, Middle Paleolithic H. sapiens juveniles show greater similarity to recent humans. These findings are consistent with recent cranial and molecular evidence for subtle developmental differences between Neanderthals and H. sapiens. When compared with earlier hominin taxa, both Neanderthals and H. sapiens have extended the duration of dental development. This period of dental immaturity is particularly prolonged in modern humans.
Early hominins formed large and thick-enamelled cheek-teeth within relatively short growth periods as compared with modern humans. To understand better the developmental basis of this process, we measured daily enamel increments, or cross striations, in 17 molars of Plio-Pleistocene hominins representing seven different species, including specimens attributed to early Homo. Our results show considerable variation across species, although all specimens conformed to the known pattern characterised by greater values in outer than inner enamel, and greater cuspal than cervical values. We then compared our results with the megadontia index, which represents tooth size in relation to body mass, for each species to assess the effect of daily growth rates on tooth size. Our results indicate that larger toothed (megadont) taxa display higher rates or faster forming enamel than smaller toothed hominins. By forming enamel quickly, large tooth crowns were able to develop within the constraints of shorter growth periods. Besides daily increments, many animals express long-period markings (striae of Retzius) in their enamel. We report periodicity values (number of cross striations between adjacent striae) in 14 new specimens of Australopithecus afarensis, Paranthropus aethiopicus, Paranthropus boisei, Homo habilis, Homo rudolfensis and Homo erectus, and show that long-period striae express a strong association with male and average male-female body mass. Our results for Plio-Pleistocene hominins show that the biological rhythms that give rise to long-period striae are encompassed within the range of variation known for modern humans, but show a lower mean and modal value of 7 days in australopithecines. In our sample of early Homo, mean and modal periodicity values were 8 days, and therefore similar to modern humans. These new data on daily rates of enamel formation and periodicity provide a better framework to interpret surface manifestations of internal growth markings on fossil hominin tooth crowns. Importantly, our data on early hominin cross striation variation may now contribute towards solving difficult taxonomic diagnoses where much may depend on fragmentary molar remains and enamel structure.
We documented the spacing and distribution of perikymata on the buccal enamel surface of fossil hominin anterior teeth with reference to a sample of modern human and modern great ape teeth. A sample of 27 anterior teeth attributed to Australopithecus (5 to A. afarensis, 22 to A. africanus) and of 33 attributed to Paranthropus (6 to P. boisei, and 27 to P. robustus) were replicated and sputter-coated with gold to enable reflected light microscopy of their surface topography. Anterior teeth were then divided into 10 equal divisions of buccal crown height. The total perikymata count in each division of crown height was recorded using a binocular microscope fitted with a vernier micrometer eyepiece. Then the mean number of perikymata per millimeter was calculated for each division. Similar comparative data for a modern sample of 115 unworn human anterior teeth and 30 African great ape anterior teeth were collected from ground sections. Perikymata counts in each taxon (together with either known or presumed periodicities of perikymata) were then used to estimate enamel formation times in each division of crown height, for all anterior tooth types combined. The distributions of these estimates of time taken to form each division of crown height follow the same trends as the actual perikymata counts and differ between taxa in the same basic way. The distinction between modern African great apes and fossil hominins is particularly clear. Finally, we calculated crown formation times for each anterior tooth type by summing cuspal and lateral enamel formation times. Estimates of average crown formation times in australopiths are shorter than those calculated for both modern human and African great ape anterior teeth. The data presented here provide a better basis for exploring differences in perikymata spacing and distribution among fossil hominins, and provide the first opportunity to describe four specimens attributed to Homo in this context. Preliminary data indicate that differences may exist among the species attributed to early Homo, especially between Homo ergaster and Homo rudolfensis on the one hand, and Homo habilis sensu strico on the other.
Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.
Most of what we know about the timing of human enamel formation comes from radiographic studies on children of known age. Here, we present new longitudinal data derived from a histological analysis of tooth enamel. Two samples, one from southern Africa and one from northern Europe, contained all anterior and molar tooth types. Two further samples contained only one tooth type: canines from a medieval Danish sample and third molars from a modern North American sample. Data were collected on 326 molars and 352 anterior teeth. Each tooth was sectioned and prepared for polarized light microscopy. We used daily enamel cross striations to determine cuspal enamel formation time, recorded the periodicity of long-period striae in the lateral enamel, and used this value to calculate enamel formation times for each decile of crown length. We present data that reveal some of the processes whereby differences in enamel formation times arise between our samples. Mean cuspal enamel formation times were similar in southern African and northern European anterior teeth, but differed in certain molar cusps. All the southern African anterior teeth completed enamel formation earlier. The greatest difference in mean chronological age at enamel completion was 5.2 vs. 6.2 years of age in lower canines. However, enamel completion times in the molar teeth showed few differences between the samples, with mean times for the longest forming cusps all falling between 3.0 years and 3.45 years. Our data suggest fewer differences between samples and smaller ranges of variation than in many radiographic studies and present a more realistic picture of worldwide variation in enamel formation times.
Histological analyses of dental development have been conducted for several decades despite few studies assessing the accuracy of such methods. Using known-period incremental features, the crown formation time and age at death of five pig-tailed macaques (Macaca nemestrina) were estimated with standard histological techniques and compared with known ages. Estimates of age at death ranged from 8.6% underestimations to 15.0% overestimations, with an average 3.5% overestimate and a 7.2% average absolute difference. Several sources of error were identified relating to preparation quality and section obliquity. These results demonstrate that histological analyses of dental development involving counts and measurements of short- and long-period incremental features may yield accurate estimates, particularly in well-prepared material. Values from oblique sections (or most naturally fractured teeth) should be regarded with caution, as obliquity leads to inflated cuspal enamel formation time and underestimated imbricational formation time. Additionally, Shellis's formula for extension rate and crown formation time estimation was tested, which significantly overestimated crown formation time due to underestimated extension rate. It is suggested that Shellis' method should not be applied to teeth with short, rapid periods of development, and further study is necessary to validate this application in other material.
Dental tissues provide important insights into aspects of hominid palaeobiology that are otherwise difficult to obtain from studies of the bony skeleton. Tooth enamel is formed by ameloblasts, which demonstrate daily secretory rhythms developing tissue-specific structures known as cross striations, and longer period markings called striae of Retzius. These enamel features were studied in the molars of two well known South African hominid species, Australopithecus africanus and Paranthropus robustus. Using newly developed portable confocal microscopy, we have obtained cross striation periodicities (number of cross striations between adjacent striae) for the largest sample of hominid teeth reported to date. These data indicate a mean periodicity of seven days in these small-bodied hominids. Important differences were observed in the inferred mechanisms of enamel development between these taxa. Ameloblasts maintain high rates of differentiation throughout cervical enamel development in P. robustus but not in A. africanus. In our sample, there were fewer lateral striae of Retzius in P. robustus than in A. africanus. In a molar of P. robustus, lateral enamel formed in a much shorter time than cuspal enamel, and the opposite was observed in two molars of A. africanus. In spite of the greater occlusal area and enamel thickness of the molars of both fossil species compared with modern humans, the total crown formation time of these three fossil molars was shorter than the corresponding tooth type in modern humans. Our results provide support for previous conclusions that molar crown formation time was short in Plio-Pleistocene hominids, and strongly suggest the presence of different mechanisms of amelogenesis, and thus tooth development, in these taxa.
Growth and development are both fundamental components of demographic structure and life history strategy. Together with information about developmental timing they ultimately contribute to a better understanding of Neanderthal extinction. Primate molar tooth development tracks the pace of life history evolution most closely, and tooth histology reveals a record of birth as well as the timing of crown and root growth. High-resolution micro-computed tomography now allows us to image complex structures and uncover subtle differences in adult tooth morphology that are determined early in embryonic development. Here we show that the timing of molar crown and root completion in Neanderthals matches those known for modern humans but that a more complex enamel-dentine junction morphology and a late peak in root extension rate sets them apart. Previous predictions about Neanderthal growth, based only on anterior tooth surfaces, were necessarily speculative. These data are the first on internal molar microstructure; they firmly place key Neanderthal life history variables within those known for modern humans.
The shape of the enamel-dentine junction (EDJ) in primate molars is regarded as a potential indicator of phylogenetic relatedness because it may be morphologically more conservative than the outer enamel surface (OES), and it may preserve vestigial features (e.g., cuspules, accessory ridges, and remnants of cingula) that are not manifest at the OES. Qualitative accounts of dentine-horn morphology occasionally appear in character analyses, but little has been done to quantify EDJ shape in a broad taxonomic sample. In this study, we examine homologous planar sections of maxillary molars to investigate whether measurements describing EDJ morphology reliably group extant anthropoid taxa, and we extend this technique to a small sample of fossil catarrhine molars to assess the utility of these measurements in the classification of fossil teeth. Although certain aspects of the EDJ are variable within a taxon, a taxon-specific cross-sectional EDJ configuration predominates. A discriminant function analysis classified extant taxa successfully, suggesting that EDJ shape may a reliable indicator of phyletic affinity. When considered in conjunction with aspects of molar morphology, such as developmental features and enamel thickness, EDJ shape may be a useful tool for the taxonomic assessment of fossil molars.
Teeth are one of the best sources of evidence for both identification and studies of demography, biological relationships and health in ancient human communities. This text introduces the complex biology of teeth and provides a practical guide to the: • excavation, cleaning, storage and recording of dental remains • identification of human teeth including those in a worn or fragmentary state • methods for studying variation in tooth morphology • study of microscopic internal and external structure of dental tissues, and methods of age-determination • estimation of age-at-death from dental development, tooth wear and dental histology • recording of dental disease in archaeological and museum collections Dental Anthropology is the text for students and researchers in anthropology and archaeology, together with others interested in dental remains from archaeological sites, museum collections or forensic cases.
Metrical methods for describing the variance in extant hominoids
are applied to phenetic groupings of later Miocene hominoids in
order to produce palaeospecies whose variance is compatible with
that seen in living hominoids.
Enamel thickness measurements are presented for samples of living
and fossil hominoids. An index of relative enamel thickness
scaled for size has been developed and this defined four categories
of relative enamel thickness metrically: Thin enamel (mean values
of relative enamel thickness between 8.90 and 11.30), intermediate/
thin enamel (mean values between 11.30 and 14.65), intermediate/
thick enamel (mean values between 14.65 and 17.25), and thick enamel
(mean values between 17.70 and 26.20). Thin enamel has been found
in Pan, Gorilla and Hylobates; intermediate/thick enamel is found
in Pongo; and thick enamel is found in Homo. Thick enamel is also
found in Sivapithecus (17.73 - 21.69).
The distribution of enamel prism packing patterns at different depths
in hominoid enamel show that Homo, Hylobates and Sivapithecus have
almost entirely Pattern 3 enamel. Pongo has an outer thickness (less
than 25%) of Pattern 1 enamel, and Pan and Gorilla have an outer (40%)
thickness of Pattern 1 enamel overlying the Pattern 3 enamel.
Pattern 3 enamel in hominoids is formed quickly (5-7μm per day)
and has well marked Hunter-Schreger bands. Pattern 1 enamel is
formed slowly (less than 2μm per day) and has no Hunter-Schreger
bands.
On the basis of these new data the commonancestral condition of
hominoid enamel has been shown to be thin enamel which formed at
the fast (Pattern 3) rate. The common ancestor of the great ape
and human clade had thick enamel which formed at the fast (Pattern 3)
rate, and this was primitively retained in the common ancestor of
the African ape and man clade and in the exclusively hominid clade.
The common ancestor of the African apes had thin enamel, a large
proportion of which (40%) formed at the slow (Pattern 1) rate.
Thick Pattern 3 enamel evolved non-adaptively through the relative
increase in dental developmental period, and dietary factors were
only of subsequent importance in the maintenance of thick enamel.
It is generally accepted that from the late Middle to the early Late Pleistocene (∼340-90 ka BP), Neanderthals were occupying Europe and Western Asia, whereas anatomically modern humans were present in the African continent. In contrast, the paucity of hominin fossil evidence from East Asia from this period impedes a complete evolutionary picture of the genus Homo, as well as assessment of the possible contribution of or interaction with Asian hominins in the evolution of Homo sapiens and Homo neanderthalensis. Here we present a comparative study of a hominin dental sample recovered from the Xujiayao site, in Northern China, attributed to the early Late Pleistocene (MIS 5 to 4). Our dental study reveals a mosaic of primitive and derived dental features for the Xujiayao hominins that can be summarized as follows: i) they are different from archaic and recent modern humans, ii) they present some features that are common but not exclusive to the Neanderthal lineage, and iii) they retain some primitive conformations classically found in East Asian Early and Middle Pleistocene hominins despite their young geological age. Thus, our study evinces the existence in China of a population of unclear taxonomic status with regard to other contemporary populations such as H. sapiens and H. neanderthalensis. The morphological and metric studies of the Xujiayao teeth expand the variability known for early Late Pleistocene hominin fossils and suggest the possibility that a primitive hominin lineage may have survived late into the Late Pleistocene in China. Am J Phys Anthropol, 2014. © 2014 Wiley Periodicals, Inc.
The study of enamel thickness has received considerable attention in regard to the taxonomic, phylogenetic and dietary assessment of human and non-human primates. Recent developments based on two-dimensional (2D) and three-dimensional (3D) digital techniques have facilitated accurate analyses, preserving the original object from invasive procedures. Various digital protocols have been proposed. These include several procedures based on manual handling of the virtual models and technical shortcomings, which prevent other scholars from confidently reproducing the entire digital protocol. There is a compelling need for standard, reproducible, and well-tailored protocols for the digital analysis of 2D and 3D dental enamel thickness. In this contribution we provide essential guidelines for the digital computation of 2D and 3D enamel thickness in hominoid molars, premolars, canines and incisors. We modify previous techniques suggested for 2D analysis and we develop a new approach for 3D analysis that can also be applied to premolars and anterior teeth. For each tooth class, the cervical line should be considered as the fundamental morphological feature both to isolate the crown from the root (for 3D analysis) and to define the direction of the cross-sections (for 2D analysis). Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.
Enamel thickness was investigated in the mesial cusp section of 167 unworn human molars by means of non-destructive micro-CT based methodology. Serial sections of the entire crown were taken at a voxel resolution of 28 microns, and the initial volume data set of each molar was standardized in orientation to obtain a vertical section that accurately contains the dentine tips of the two mesial cusps. Enamel thickness at the cusp tips, occlusal basin, and lateral crown face was measured in the mesial cusp section and in sections offset from that section by 0.6 mm. We found that thickness at the cusp tips may be overestimated in offset sections by up to about I mm, and those of the occlusal basin overestimated or underestimated by up to about 0.5 mm. We also found that maximum 'radial' thickness of the lateral crown face was least affected by section position, usually with discrepancies of less than about 0.1-0.2 mm. In all serial positions in both upper and lower molars, a 'functional' (lingual in uppers and buccal in lowers) to 'non-functional' side gradient in enamel thickness was observed in cusp tip, occlusal basin, and lateral crown face enamel, with the exception of the characteristically thin enamel at the protoconid and paracone cusp tips. Serial differences in thickness were seen between the thinner M1 and the two posterior molars in many but not all measures of thickness, the pattern of which appears to be influenced by the thin M I mesiobuccal cusp enamel. Individual variation of maximum lateral thickness, the least variable measure of thickness, was found to be substantial (a 30-60% range) even with serial and buccolingual positions controlled. Correlation between whole crown average enamel thickness and maximum lateral thickness was high, indicating that the latter is a potentially useful predictor of overall enamel thickness of the molar crown. The present results indicate that interspecific comparisons of enamel thickness must be made with careful attention to positional placement of thickness measures, potential serial differences, and intraspecific variation.
Molar enamel thickness is a key feature in the study of hominid evolution. Our understanding of enamel thickness and distribution patterns, however, has so far been based mostly on the limited information available from physical cross sections of the crown. In this study, the 3-dimensional (3D) whole crown enamel distribution pattern was explored in 74 extant great ape and modem human molars. Serial cross sections obtained from microfocal X-ray computed tomography were used to generate digital molar reconstructions at 50 to 80 micron voxel resolution, each crown represented by two to five million voxels. Surface data of both enamel dentine junction (EDJ) and outer enamel were extracted to derive volumetric measures, surface areas, curvilinear distances, and whole crown radial thickness maps. Three-dimensional average enamel thickness (AET) was defined as enamel volume divided by EDJ surface area. In 3D AET relative to tooth size, Homo exhibited the thickest, Gorilla the thinnest, and Pan and Pongo intermediately thick enamel. This result differs from previous claims that molar enamel of Pongo is relatively thicker than that of Pan. The discrepancy between three and two-dimensional (2D) values of AET stems from a combination of local differences in within tooth enamel distribution pattern and EDJ topography between Pan and Pongo molars. It demonstrates that 2D AET is not an appropriate substitute or estimator of whole crown AET. Examination of whole crown 3D distributions of molar enamel revealed a pattern common to all four examined species, the "functional" side of the molar having thicker enamel than the opposite side. However, some unique aspects of each species were also apparent. While the Gorilla molar has relatively thin enamel throughout its crown, Pan molars are characterized by particularly thin enamel in the occlusal fovea, and Pongo molars by an accentuation of relatively thin basal and thick occlusal enamel. Human molars are characterized by relatively thick enamel throughout the crown, with relatively large contrasts between buccal and lingual, and between mesial and distal crown portions. The ancestral condition common to the four extant species can be estimated by interpreting molar enamel distribution patterns unique to each genus as likely to be derived. We hypothesize that the last common ancestor likely had intermediately thick enamel, without particular thickening (or thinning) of enamel either occlusally or basally.
The excavations carried out in the cave of Santa Ana (Cáceres, Spain), cave of the karstic network of the “Calerizo” of Cáceres, enabled us to know the existence, in stratigraphy, of the three lito-techniques modes which characterize the industrial development of lower and middle Pleistocene in the Iberian Peninsula. On standby of new research, the results obtained until now permit us two work out a diachronic assumption of technical evolution. In the Iberian Peninsula, there are only two karstic systems where we can fallow this technological development; one is the “Sierra de Atapuerca” (Burgos) and the other is the “Calerizo de Cáceres” (Cáceres).
The formation of lateral enamel in Neandertal anterior teeth has been the subject of recent studies. When compared to the anterior teeth of modern humans from diverse regions (Point Hope, Alaska; Newcastle upon Tyne, England; southern Africa), Neandertal anterior teeth appear to fall within the modern human range of variation for lateral enamel formation time. However, the lateral enamel growth curves of Neandertals are more linear than those of these modern human samples. Other researchers have found that the lateral enamel growth curves of Neandertals are more linear than those of Upper Paleolithic and Mesolithic modern humans as well. The statistical significance of this apparent difference between Neandertal and modern human lateral enamel growth curves is analyzed here. The more linear Neandertal enamel growth curves result from the smaller percentage of total perikymata located in the cervical halves of their teeth. The percentage of total perikymata in the cervical halves of teeth is therefore compared between the Neandertal sample (n = 56 teeth) and each modern human population sample: Inuit (n = 65 teeth), southern African (n = 114 teeth), and northern European (n = 115 teeth). There are 18 such comparisons (6 tooth types, Neandertals vs. each of the three modern human populations). Eighteen additional comparisons are made among the modern human population samples. Statistically significant differences are found for 16 of the 18 Neandertal vs. modern human comparisons but for only two of the 18 modern human comparisons. Statistical analyses repeated for subsamples of less worn teeth show a similar pattern. Because surface curvature is thought to affect perikymata spacing, we also conducted measurements to assess surface curvature in thirty teeth. Our analysis shows that surface curvature is not a factor in this lateral enamel growth difference between Neandertals and modern humans.
The systematic excavation of the Sima de los Huesos (SH) site in Sierra de Atapuerca (Burgos, Spain) has yielded the largest hominin collection worldwide for the Middle Pleistocene. The dental sample now consists of more than 500 teeth that provide exceptional opportunities to define the dental morphological pattern of a Middle Pleistocene population as well as develop hypotheses about the origins of the Neanderthals. The dental collection has now increased to over 533 specimens (525 permanent and 8 deciduous teeth), necessitating new morphological assessments. Thus, we present a detailed morphological description of the SH permanent dentition recovered up to 2007, accomplishing comparisons with European Middle Pleistocene hominins, Neanderthals, and early and contemporary Homo sapiens. We find that SH dentitions present all the morphological traits that, either in their degree of expression, frequency, or particular combination, are usually considered as typical of Homo neanderthalensis. This study ratifies the deep roots of the Neanderthal lineage in the Middle Pleistocene of Europe. In addition, SH teeth are morphologically "more Neanderthal" than other penecontemporaneous Middle Pleistocene samples such as Mauer or Arago, and even more derived than some classic Neanderthal samples. Thus, our study would not sustain the linearity of the accretion process hypothesized for the origins of the Neanderthals, and we suggest that other evolutionary models and scenarios should be explored for the Middle and Upper Pleistocene of Europe. We propose that more than one hominin lineage may have coexisted during the Middle Pleistocene in Europe.
This paper continues the series of articles initiated in 2006 that analyse hominin dental crown morphology by means of geometric morphometric techniques. The detailed study of both upper premolar occlusal morphologies in a comprehensive sample of hominin fossils, including those coming from the Gran Dolina-TD6 and Sima de los Huesos sites from Atapuerca, Spain, complement previous works on lower first and second premolars and upper first molars. A morphological gradient consisting of the change from asymmetric to symmetric upper premolars and a marked reduction of the lingual cusp in recent Homo species has been observed in both premolars. Although percentages of correct classification based on upper premolar morphologies are not very high, significant morphological differences between Neanderthals (and European middle Pleistocene fossils) and modern humans have been identified, especially in upper second premolars. The study of morphological integration between premolar morphologies reveals significant correlations that are weaker between upper premolars than between lower ones and significant correlations between antagonists. These results have important implications for understanding the genetic and functional factors underlying dental phenotypic variation and covariation.
This article is the third of a series that explores hominin dental crown morphology by means of geometric morphometrics. After the analysis of the lower second premolar and the upper first molar crown shapes, we apply the same technique to lower first premolar morphology. Our results show a clear distinction between the morphology seen in earlier hominin taxa such as Australopithecus and African early Homo, as well as Asian H. erectus, and more recent groups such as European H. heidelbergensis, H. neanderthalensis, and H. sapiens. The morphology of the earlier hominins includes an asymmetrical outline, a conspicuous talonid, and an occlusal polygon that tends to be large. The morphology of the recent hominins includes a symmetrical outline and a reduced or absent talonid. Within this later group, premolars belonging to H. heidelbergensis and H. neanderthalensis tend to possess a small and mesiolingually-displaced occlusal polygon, whereas H. sapiens specimens usually present expanded and centered occlusal polygons in an almost circular outline. The morphological differences among Paranthropus, Australopithecus, and African early Homo as studied here are small and evolutionarily less significant compared to the differences between the earlier and later homin taxa. In contrast to the lower second premolar and the upper first molar crown, the inclusion of a larger hominin sample of lower first premolars reveals a large allometric component.
A recent evaluation of upper first molar (M¹) crown size and cusp proportions in the genus Homo (Quam et al. 2009) describes Homo antecessor as maintaining a primitive pattern of cusp proportions, similar to that identified in australopithecines and the earliest members of the genus Homo. These results contrast with those of Gómez-Robles et al. (2007), who described the crown shape in these molars as derived and similar to Neanderthals and European Homo heidelbergensis. The reassessment of these measurements following the same methodology described by Quam et al. (2009) in all the M(1) s that are currently part of the hypodigm of H. antecessor demonstrates that the fossils from TD6 not only have the same cusp proportions identified in later Homo species, but also a strongly reduced metacone and a large hypocone shared with Middle and Upper Pleistocene members of the Neanderthal lineage. The evolutionary significance of these features should be evaluated in light of the results provided by recently discovered dental, cranial, mandibular, and postcranial H. antecessor fossils.
The aim of this study was to describe similarities and differences in the way modern and fossil hominin teeth grow in height. Measurements from longitudinal ground sections of 7 modern human canines and 19 first permanent molars were used to calculate extension rates in the crowns and roots and to plot distance curves for growth in tooth height. These were compared with identical data for 3 fossil hominin teeth attributed respectively to Paranthropus robustus, Homo erectus and Homo neanderthalensis. Enamel extension rates in each of the three fossil taxa fell within the range of modern humans. Root extension rates in the fossil taxa also fell within modern human ranges but differed in their pattern with either an early or late marked increase in root length. Extension rates in the canine crowns were higher in cuspal enamel than in lateral enamel. Combinations of high or low cuspal enamel extension rates, with either longer or shorter times taken to form lateral enamel, explain how crown formation times may vary independently of completed crown heights.
Enamel thickness has assumed unique importance in the debate about the hominid status of Ramapithecus, despite the fact that there is little agreement about the meaning of 'thick enamel' or the significance of enamel thickness for hominoid taxonomy. My aim here is to evaluate the usefulness of enamel thickness and microstructure as characteristics for determining the relationships of the later Miocene hominoids, based both on a quantitative study of enamel thickness in extant hominoids and four species of later Miocene Sivapithecus (including 'Ramapithecus') and on scanning electron microscope analysis of enamel microstructure. Four categories of enamel thickness are defined metrically here and have been related to enamel microstructure and developmental rates. Thin fast-formed (pattern 3) enamel represents the ancestral condition in hominoids; it increased developmentally to thick pattern 3 enamel in the great ape and human clade and that condition is primitively retained in Homo and in the fossil hominoid Sivapithecus (including 'Ramapithecus'). Enamel thickness has been secondarily reduced in the African apes and also, although at a different rate and extent, in the orang-utan. Thick enamel, previously the most important characteristic in arguments about the earliest hominid, does not therefore identify a hominid.
Among primitive peoples dental attrition appears to be a natural phenomenon. Often the degrees and kinds of tooth wear vary from population to population. This variability is possibly related to certain material aspects of culture such as diet, food preparation techniques and tool usage. In order to learn more about these relationships, extensive cross cultural comparisons must be made.
This paper reports on a study of dental attrition among skeletal remains of North American Indians from three areas: California, the Southwest and the Valley of Mexico. A method of comparing worn teeth of these populations was devised so several characteristics of the teeth and supporting bone could be examined by population. This study showed significant differences in type and degree of wear among the three groups as well as differences between sexes within each population.
A positive correlation between tooth wear and cultural factors was found. Dietary specialization and division of labor appear to be responsible for the degree and type of wear found in this sample. Further studies of this type are planned to expand the sample size and, if the new data support these correlations, valuable information about human–environmental relationships can be gained.
This study presents data on the enamel thickness of deciduous (dm2) and permanent (M1-M3) molars for a geographically diverse sample of modern humans. Measurements were recorded from sections through the mesial cusps of unworn teeth. Enamel is significantly thinner on deciduous than on permanent molars, and there is a distinct trend for enamel to increase in relative thickness from M1 to M3. The relatively thicker enamel of M2s and especially M3s can be related to the overall reduction in size of more distal molar crowns, which has been attained through a differential loss of the dentine component. Enamel tends to be thicker on the protocone than on the paracone, and thicker on the protoconid than on the metaconid, but its distribution is not wholly concordant with models that predict increased thickness as a means by which to counter heavier attritional loss on these "functional" cusps. Indeed, the thickness of enamel tends to be more variable on cusp tips and occlusal surfaces than over the lateral aspects of cusps. The proportionately thicker enamel over the lateral aspects of the protocone and protoconid more likely serves as a means to prolong functional crown life by preventing cusp fracture, rather than being an adaptation to increase the attritional longevity of wear facets. The present data suggest that the human dentition is not predisposed to develop a helicoidal wear plane through the disposition of molar enamel thickness.
Tooth enamel thickness has long been an important character in studies of primate and especially hominin phylogeny, taxonomy, and adaptation. Current methods for accurately assessing enamel thickness involve the physical sectioning of teeth, because measurements of enamel thickness using some radiographic techniques are unreliable. However, because destructive methods limit sample sizes and access to important fossil specimens, it is desirable that they be replaced with nondestructive techniques. Although microfocal X-ray computed tomography (mCT) has been used recently in studies of enamel thickness, the accuracy of this technique has yet to be established. The present research compares physical sections to computer-generated mCT sections of teeth from a variety of primate and nonprimate, recent and fossil taxa to examine whether enamel thickness, tooth size, and diagenetic remineralization (fossilization) impact the ability of mCT to measure enamel thickness accurately. Results indicate that recent teeth of varying size and thickness are clearly and accurately depicted in mCT scans, with measurements from nearly identical planes in physical and mCT sections differing by 3-5%. A fossil papionin molar (ca. 2 Myr) was also accurately measured using mCT scans, although thinner enamel in much older therapsid (ca. 263-241 Myr) teeth could not be distinguished from dentine. mCT is thus an accurate technique for measuring enamel thickness in recent taxa, although heavily mineralized teeth pose an obstacle to the ability of mCT to distinguish dental tissues. Moreover, absolutely thin enamel (less than approximately 0.10 mm) is difficult to resolve adequately in raw mCT images based on pixel values alone. Therefore, caution must be exercised in the application of mCT to the study of fossilized teeth.
Mandibular premolars are increasingly used in taxon-specific diagnostic analyses of hominins. Among the principal difficulties in these evaluations is the absence of discrete, discernible, and comparable anatomical structures for rigorous quantitative assessment. Previous research has addressed either internal crown surface features (such as cusps and fossae) or the morphology of the crown outline. In the present paper, we integrate both types of information in the examination of morphological variation of lower P4s (n = 96) among various fossil hominin species with an emphasis on genus Homo. We use a set of 34 2D landmarks combining coordinate data from four classical dental landmarks on the occlusal surface and 30 sliding semilandmarks of the crown outline. Our results indicate that external shape variation is closely related to the configuration of the occlusal morphological features and influenced by dental size. The external and internal shapes of P4 are polymorphic but still useful in depicting a primitive-derived gradient. The primitive pattern seems to have been an asymmetrical contour with a mesially displaced metaconid, development of a bulging talonid, and a broad occlusal polygon. The trend toward dental reduction during the Pleistocene produced different morphological variants with a reduced occlusal polygon and decreased lingual occlusal surface in later Homo species. Homo heidelbergensis/neanderthalensis have fixed plesiomorphic traits in high percentages, whereas in modern humans a symmetrical outline with a centered metaconid and talonid reduction evolved.
Enamel is formed incrementally by the secretory activity of ameloblast cells. Variable stages of secretion result in the formation of structures known as cross striations along enamel prisms, for which experimental data demonstrate a correspondence with daily periods of secretion. Patterns of variation in this daily growth are important to understanding mechanisms of tooth formation and the development of enamel thickness. Transmitted light microscopy (TLM) of histological ground sections and scanning electron microscopy (SEM) of bulk specimens or their surface replicas are the usual methods for investigating cross striations. However, these methods pose some constraints on the study of these features in Plio-Pleistocene hominid enamel, the specimens of which may only rarely be sectioned for TLM or examined on only their most superficial surfaces for SEM. The recent development of portable confocal scanning optical microscopy (PCSOM) resolves some of the restrictions on fractured enamel surfaces, allowing the visualization of cross striations by direct examination. This technology has been applied here to the study of Australopithecus africanus and Paranthropus robustus hominid molars from the Plio-Pleistocene of South Africa. We hypothesize that these taxa have increased enamel appositional rates compared with modern humans, because despite having thicker enamelled molars (particularly P. robustus), the enamel crowns of these fossil taxa take an equivalent or reduced amount of time to form. Cross striations were measured in cuspal, lateral and cervical regions of the enamel crowns, and, within each region, the inner, middle and outer zones. Values obtained for A. africanus outer zones of the enamel crown are, in general, lower than those for P. robustus, indicating faster forming enamel in the latter, while both taxa show higher rates of enamel growth than modern humans and the African great apes. This demonstrates a relatively high degree of variability in the mechanisms underlying the development of enamel across taxa.
The Plio-Pleistocene site of Kromdraai, South Africa, is well known for the recovery of the holotype of Paranthropus robustus, one of nine individual hominids recovered from this site to date. Among the Kromdraai sample, the specimen KB 5223 comprises several isolated deciduous and permanent lower teeth assigned to Paranthropus, the only recognized genus at this site. However, a more recent analysis of this specimen suggested that it should be classified as Homo. The lower right first permanent molar of KB 5223 had been previously sectioned along the tips of the mesial cusps, exposing its enamel microstructure. Previous studies had indicated differences between Homo and Paranthropus at the microstructural level. A portable confocal scanning microscope was used to describe details of the enamel microstructure of the M1 and I1 of this specimen. Angles formed between the striae of Retzius and the enamel dentine junction (EDJ), daily secretion rates in cuspal enamel of the protoconid and metaconid and crown formation time of the RM1 are provided. The number of perikymata on the right I1 was counted. Results indicate that some features recorded in the KB 5223 molar differ from those of Paranthropus. However, the number of perikymata on the I1 is lower than values so far reported for early Homo but similar to Paranthropus. Crown formation time of KB 5223 M1 was markedly lower than mean values of M1 in H. sapiens, but similar to other early hominids. Daily secretion rates in the cuspal enamel of KB 5223 M1 were higher than in modern humans.
Methods of measuring tissue area from images of longitudinal thin tooth sections have been used to assess sexual dimorphism in the permanent dentition. The aim of this study was to demonstrate the extent of sexual dimorphism within the coronal tissue proportions of permanent mandibular canines and premolars, using area measurements of the enamel and dentine-pulp core. The sample consisted of embedded "half-tooth" sections from 45 individuals, all of known age-at-death and sex, collected from the St. Thomas' Anglican Church historic (1821-1874) cemetery site in Belleville, ON, Canada. The relative dentine-pulp area of the third premolars and canines displayed high levels of sexual dimorphism, as well as statistically significant mean differences between the sexes. The male canines and premolars have significantly more dentine than their female counterparts, as well as relatively more dentine with respect to overall crown size. The female canines and premolars have significantly more enamel relative to overall crown area than those of the males. These results suggest that relative area measures of crown tissues are more predictable measures of sexual dimorphism than absolute measures, and tissue proportions may remain constant despite intrasex variation in overall tooth crown size.
Molar enamel thickness has played an important role in the taxonomic, phylogenetic, and dietary assessments of fossil primate teeth for nearly 90 years. Despite the frequency with which enamel thickness is discussed in paleoanthropological discourse, methods used to attain information about enamel thickness are destructive and record information from only a single plane of section. Such semidestructive planar methods limit sample sizes and ignore dimensional data that may be culled from the entire length of a tooth. In light of recently developed techniques to investigate enamel thickness in 3D and the frequent use of enamel thickness in dietary and phylogenetic interpretations of living and fossil primates, the study presented here aims to produce and make available to other researchers a database of 3D enamel thickness measurements of primate molars (n=182 molars). The 3D enamel thickness measurements reported here generally agree with 2D studies. Hominoids show a broad range of relative enamel thicknesses, and cercopithecoids have relatively thicker enamel than ceboids, which in turn have relatively thicker enamel than strepsirrhine primates, on average. Past studies performed using 2D sections appear to have accurately diagnosed the 3D relative enamel thickness condition in great apes and humans: Gorilla has the relatively thinnest enamel, Pan has relatively thinner enamel than Pongo, and Homo has the relatively thickest enamel. Although the data set presented here has some taxonomic gaps, it may serve as a useful reference for researchers investigating enamel thickness in fossil taxa and studies of primate gnathic biology.
Two hypotheses, based on previous work on Neandertal anterior and premolar teeth, are investigated here: (1) that estimated molar lateral enamel formation times in Neandertals are likely to fall within the range of modern human population variation, and (2) that perikymata (lateral enamel growth increments) are distributed across cervical and occlusal halves of the crown differently in Neandertals than they are in modern humans. To investigate these hypotheses, total perikymata numbers and the distribution of perikymata across deciles of crown height were compared for Neandertal, northern European, and southern African upper molar mesiobuccal (mb) cusps, lower molar mesiobuccal cusps, and the lower first molar distobuccal (db) cusp. Sample sizes range from five (Neandertal M(1)db) to 29 (southern African M(1)mb). Neandertal mean perikymata numbers were found to differ significantly from those of both modern human samples (with the Neandertal mean higher) only for the M(2)mb. Regression analysis suggests that, with the exception of the M(2)mb, the hypothesis of equivalence between Neandertal and modern human lateral enamel formation time cannot be rejected. For the M(2)mb, regression analysis strongly suggests that this cusp took longer to form in the Neandertal sample than it did in the southern African sample. Plots of perikymata numbers across deciles of crown height demonstrate that Neandertal perikymata are distributed more evenly across the cervical and occlusal halves of molar crowns than they are in the modern human samples. These results are integrated into a discussion of Neandertal and modern human lateral enamel formation across the dentition, with reference to issues of life history and enamel growth processes.
A recent study demonstrated that variation in enamel cap crown formation in the anterior teeth is greater than that in the molars from two geographically distinct populations: native indigenous southern Africans and northern Europeans. Eighty southern African and 69 northern European premolars (P3 and P4) were analyzed in the present study. Cuspal, lateral, and total enamel formation times were assessed. Although cuspal enamel formation times were not consistently different between the two populations, both lateral and total enamel formation times generally were. Bonferroni-corrected t-tests showed that southern Africans had significantly shorter lateral enamel formation time for five of the six cusps, as well as significantly shorter total enamel formation time for these same cusps. An analysis of covariance performed on the lingual cusps of the upper third and fourth premolars showed that differences in enamel formation times between these populations remained when crown height was statistically controlled. A further goal of this study was to ascertain, based on perikymata counts, what Neandertal periodicities would have to be in order for their teeth to have lateral enamel formation times equivalent to either southern Africans or northern Europeans. To this end, perikymata were counted on 32 Neandertal premolars, and the counts were inserted into regression formulae relating perikymata counts to periodicity for each population and each tooth type. Neandertal enamel formation times could be equivalent to those of southern Africans or northern Europeans only if their hypothetical periodicities fall within the range of periodicities for African apes and modern humans (i.e., 6-12 days). The analysis revealed that both populations could encompass Neandertal timings, with hypothetical periodicities based on the southern African population necessitating a lower range of periodicity (6-8 days) than those based on the northern European population (8-11 days).
The timing and sequence of enamel development, as well as enamel thickness, was documented for individual cusps (protoconid, hypoconid, metaconid, entoconid) in 15 unworn permanent lower first molars (M(1)s) from a sample of modern human juveniles. These data were compared with previously published data for modern and fossil species reported in the literature. Crown formation in all teeth was initiated in the protoconid and completed in the hypoconid. These cusps had significantly longer formation times (2.91 and 2.96 yrs, respectively) than the metaconid and entoconid (2.52 and 2.38 yrs, respectively), as well as thicker enamel, and each represented between 92-95% of the total crown formation time. Rates of enamel secretion in all cusps increased significantly from 2.97 microm in the inner enamel to 4.47 microm in the outer enamel. Two cusps of one individual were studied in more detail and did not follow this typical trajectory. Rather, there was a sharp decrease in the middle of enamel formation and then a slow recovery of secretion rates from the mid- to outer enamel. This anomalous trajectory of enamel formation is discussed in the context of other nondental tissue responses to illness. Neither secretion rates nor periodicity differed significantly when compared between the cusps of each molar. Differences in cusp formation times, initiation, and completion suggest a relationship between the rates of enamel formation and enamel thickness. This fits with expectations about the mechanics of the chewing cycle and general lower molar morphology. A comparison with similar data for some nonhuman primates and fossil hominoids suggests this relationship may hold true across several primate taxa. Other aspects of enamel growth differed between this human sample and certain fossil species. The lower molars formed slowly over a longer period of time, which may reflect the extended growth period of modern humans. The methodological approach adopted in this study is discussed in the context of that used in other studies.
Effects of molar crown orientation to measures of lateral enamel thickness in the mesial cusp section
- Kono
Kono, R. T., & Suwa, G. (2005). Effects of molar crown orientation to
measures of lateral enamel thickness in the mesial cusp section. Bulletin of the National Science Museum. Series B, 31, 11-22.
Cuevas para la eternidad: sepulcros prehistóricos de la provincia de cáceres
- C. E. Cerrillo
- C. A. González