Sexual dimorphism in crown units of mandibular deciduous and permanent molars in Australian Aborigines.

Sexual differences in the crown units of mandibular molars were investigated in Australian Aborigines. The first and second deciduous molars (dm1 and dm2), and first to third permanent molars (M1, M2 and M3) were measured on dental casts using a sliding caliper. Measurements of tooth crowns included overall mesiodistal and buccolingual diameters, as well as the mesiodistal and buccolingual diameters of the trigonid and talonid. Percentage dimorphism values were greater in the talonid dimensions than the trigonid, indicating that sex differences tend to be larger in the later-developing crown units. Sex differences in mesiodistal diameters increased from dm1 to M2 but decreased for M3, the tooth that showed the least dimorphism of all the molars. This result seems to be due to the marked variability in size of the M3 between individuals.     

Carabelli's trait and tooth size of human maxillary first molars

Carabelli's trait is a morphological feature that can occur on the protocone of human maxillary molars. This study tests the hypothesis that Carabelli's trait is correlated statistically with the dimensions of the crown's four principal cusps or whether, as a cingular feature, the trait truly accretes onto an otherwise unaffected crown. Computer-assisted image analysis was used to measure the 6 intercusp distances and 12 angular relationships among cusp tips on the permanent first molar of 300 young adult American whites. Carabelli's complex was scored using an 8-grade ordinal scheme. Crown size was quantified in three ways, namely as 1) maximum mesiodistal and buccolingual diameters, 2) the 6 intercusp distances, and 3) the 12 angular cusp arrangements. There was no sex difference in the morphological expression of Carabelli's trait in this sample. Overall crown size and intercusp distances were significantly and progressively larger in molars with larger Carabelli's trait expressions. There are graded size responses between crown size (mesiodistal and buccolingual diameters), sizes of the four principal cusps, and morphological stage of Carabelli's complex, though the statistical relationships are appreciably stronger in males than females. Carabelli's trait occurs preferentially in larger molars. In contrast, angular (shape) relationships among cusp tips are not discernibly affected by trait size in either sex. There is the situation, then, that Carabelli's trait is developmentally correlated with crown size, but with no apparent alteration of cusp arrangements, suggesting that the increases are isometric across the occlusal table. Why the association is much weaker in females remains speculative, but these data provide yet another line of evidence that, within a population, tooth size is associated in a positive fashion with crown complexity.     

A principal components analysis of human odontometrics

It has long been recognized that tooth crown diameters in hominoids are all positively intercorrelated one with another. This study reports on sex-specific correlation matrices derived from 2,650 individuals from the Solomon Islands, Melanesia. Mesiodistal and buccolingual diameters of all permanent teeth from one side are used, excluding third molars. Analysis discloses significant sex dimorphism in the strengths of the intercorrelations, with females being better integrated. Principal components analysis (PCA) provides an objective means of data reduction (shown here to be preferable to simple size summation methods) and decorrelation of the resulting linear combinations. Four components are extracted (with results being virtually identical in the two sexes) and arguments are put forth that varimax rotation to "a simpler solution" may be counterproductive. Before rotation, the four components are 1) overall size, 2) buccolingual widths contrasted with mesiodistal lengths, 3) anterior (I,C) contrasted with posterior (P,M) teeth, and 4) premolars contrasted with molars. Most of the explained (shared) variance (63%) extracted by PCA is in overall size of the dentition. There is a strong urge to view the results of these principal components analyses as reflective of biologically and genetically meaningful entities.     

Intercusp relationships of the permanent maxillary first and second molars in American whites

Much of a human molar's morphology is concentrated on its occlusal surface. In view of embryologists' recent attention on the determination of crown morphology by enamel knots that initiate cusp formation, we were interested in the arrangement of cusp apices in the definitive tooth. Computer-assisted image analysis was used to measure intercusp distances and angles on permanent maxillary M1 and M2 in a sample of 160 contemporary North American whites. The intent was to generate normative data and to compare the size and variability gradients from M1 to M2. There is little sexual dimorphism in intercusp distances or angles, even though the conventional mesiodistal (MD) and buccolingual (BL) crown size is 2.0% and 4.0% larger in males, respectively, in these same teeth. Dimensions decreased in size and increased in variability from M1 to M2, but differentially. Cusps of the trigon were more stable between teeth, especially the paracone-protocone relationship. Principal components analysis on the six M1 distances disclosed only one eigenvalue above 1.0, indicating that overall crown size itself is the paramount controlling factor in this tooth that almost invariably exhibits a hypocone. In contrast, four components were extracted from among the 12 angular cusp relationships in M1. These axes of variation may prove useful in studies of intergroup differences. A shape difference occurs in M2, depending on whether the hypocone is present; when absent, the metacone is moved lingually, creating more of an isosceles arrangement for the cusps of the trigon. Statistically, correlations are low between occlusal intercusp relationships and conventional crown diameters measured at the margins of the crowns that form later. Weak statistical dependence between cusp relationships and traditional MD and BL diameters suggest that separate stage- and location-specific molecular signals control these different parts (and different stages) of crown formation.     

Alternative dental measurements: proposals and relationships with other measurements

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

Development and size of the teeth of Macaca mulatta.

A cross-sectional sample of 151 skulls from Macaca mulatta of known age and similar rearing in U.S. Primate Centers was analyzed to determine age-related "norms" of stages of development and size of teeth. The stages of development from the follicle of a deciduous incisor in the fetus to completion of the root with apex closed of the permanent third molar were related to age. The age range observed for eruption of each tooth was noted and related to its stage of development. The crown of each erupted tooth was found to be completely developed, but growth of its root continued for a longer, indeterminate period. When a deciduous tooth was exfoliated, the crown of the permanent successor was found to be completed and root growth had begun. Measurements of both mesiodistal and faciolingual diameters and of crown length of the teeth in situ and of total length and root length on roentgenograms were examined for sexual dimorphism. The faciolingual diameter of the deciduous mandibular second incisor and of both second molars showed the greatest sexual dimorphism among both diameters of all deciduous teeth. The mesiodistal and faciolingual diameters of the mandibular premolars were found to be the best dimensions in discriminant functions for identifying sex in the absence of permanent canines.     

Tissue proportions and enamel thickness distribution in the early Middle Pleistocene human deciduous molars from Tighenif,Algeria

The present study of three human upper deciduous molars from the early Middle Pleistocene site of Tighenif, Algeria, constitutes the first microtomographic-based endostructural exploration of African fossil teeth likely representative of the Homo heidelbergensis morph. Comparative morphological observations and 2-3D measurements describing subtle tooth organization (crown tissue proportions) and enamel thickness topography (site-specific distribution and global patterning) indicate that their virtual extracted structural signature better fits the modern human, rather than the Neanderthal condition. Accordingly, we predict that the inner structural morphology of the deciduous molars from the Middle Pleistocene western European series better fits the primitive, and not the derived Neanderthal figures.     

Tissue contributions to sex and race: differences in tooth crown size of deciduous molars.

This study describes size of constituent deciduous tooth crown components (enamel, dentine, and pulp) to address the manner in which males characteristically have larger teeth than females, and the observation that teeth of American blacks are larger than those of American whites. Measurements were collected (n = 333 individuals) from bitewing radiographs using computer-aided image analysis. Tissue thicknesses (enamel, dentine, pulp) were measured at the crown's mesial and distal heights of contour. Deciduous mesiodistal molar crown length is composed of about 1/7 enamel, 1/3 dentine, and 1/2 pulp. Details differ by tooth type, but males typically have significantly larger dentine and pulp dimensions than females; there is no sexual dimorphism in marginal enamel thickness. Males scale isometrically with females for all variables tested here. Blacks significantly exceed whites in size of all tissues, but tissue types scale isometrically with blacks and whites with one exception: enamel thickness is disproportionately thick in blacks. While the absolute difference is small (5.56 mm of enamel in blacks summed over all four deciduous molar tooth types vs. 5.04 mm in whites), the statistical difference is considerable (P < 0.001). Aside from enamel, crown size in blacks is increased proportionately vis-à-vis whites. Principal components analysis confirmed these univariate relationships and emphasizes the statistical independence of crown component thicknesses, which is in keeping with the sequential growth and separate embryonic origins of the tissues contributing to a tooth crown. Results direct attention to the rates of enamel and dentine deposition (of which little is known), since the literature suggests that blacks (with larger crowns and thicker enamel) spend less time in tooth formation than whites.     

Additional human remains from Blombos Cave, South Africa: (1999-2000 excavations)

The uppermost Middle Stone Age (MSA) layers at Blombos Cave contain high densities of Still Bay bifacial points. Information from other regional sites places the Still Bay prior to the Howiesons Poort industry, which has been dated at 65-70 ka. The Blombos Cave MSA strata have yielded nine human teeth or dental fragments. Four that were recovered during the 1997-1998 excavations have been published elsewhere. The remaining five were discovered during the 1999-2000 field seasons; these are described here. Three of the new specimens are deciduous teeth, and two are permanent premolar and molar crown fragments. The entire dental sample probably represents at least five and as many as seven individuals. The deciduous teeth from the upper MSA levels are likely to have been exfoliated in the cave. One deciduous tooth and the permanent tooth fragments from the lower MSA levels probably represent three individuals who died in or near the cave. The Blombos Cave premolars preserve horizontal circum-cervical striae suggestive of palliative tooth pick use. Approximately half of the permanent and deciduous crown diameters exceed those of recent Africans; for the remainder, the fossil values fall among modern African sample means. The Blombos Cave tooth crowns tend to be smaller than the majority of penecontemporaneous Neandertal teeth. The morphology of the Blombos Cave di is comparable to MSA homologues from the nearby, and presumably somewhat younger site of Die Kelders Cave 1.     

Relationships between age and dental attrition in Australian aboriginals

Tooth wear scores (ratios of exposed dentin to total crown area) were calculated from dental casts of Australian Aboriginal subjects of known age from three populations. Linear regression equations relating attrition scores to age were derived. The slope of the regression line reflects the rate of tooth wear, and the intercept is related to the timing of first exposure of dentin. Differences in morphology between anterior and posterior teeth are reflected in a linear relationship between attrition scores and age for anterior teeth but a logarithmic relationship for posterior teeth. Correlations between age and attrition range from less than 0.40 for third molars (where differences in the eruption and occlusion of the teeth resulted in different patterns of wear) to greater than 0.80 for the premolars and first molars. Because of the generally high correlations between age and attrition, it is possible to estimate age from the extent of tooth wear with confidence limits of the order of +/- 10 years.     

Variation in modern human premolar enamel formation times: implications for Neandertals

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).     

Secular trend in tooth size in urban Chinese assessed from two-generation family data

Environmental improvements can lead to greater size of skeletodental structures as the population comes closer to expressing its genetic growth potential. Such secular trends have been documented in many human settings, especially for increased stature and faster tempos of growth. The present study is based on 185 same-sex parent-offspring pairs of data for maximum buccolingual crown diameters of the permanent teeth from a cohort in Beijing, China, where parents experienced much of their development during and after World War II with the deprivations of the changing regime. Their offspring enjoyed the relative stability of the established Communist system, where nutrition and stability were much improved. There were significant increases in buccolingual diameters of the premolars and molars in the offspring. Increase for premolars and molars was about 1%, but larger in females than males (1.6% vs. 0.5%). Opposite changes occurred in the incisors and canines (i.e., larger parental dimensions), but we contend that these are an age-related artifact brought on by greater passive eruption of older individuals' teeth that exposes a broader portion of the crown at the gingival margin. The secular trend in crown size coincides with other research in contemporary China, disclosing increases in body size and faster tempos of growth as health and nutrition continue to improve.     

A new spalacolestine mammal from the Early Cretaceous Jehol Biota and implications for the morphology,phylogeny, and palaeobiology of Laurasian ‘symmetrodontans’

‘Symmetrodontans’ are extinct mammals characterized by having a reversed‐triangle molar pattern in which three main cusps define a triangular molar crown. This dental morpholgy has been regarded as being intermediate between the ‘triconodont’ tooth and the tribosphenic pattern characterizing therians; it is a key feature in taxonomy of Mesozoic mammals and one to understand mammalian evolution and palaeobiology. Here we report a new genus and species of ‘symmetrodontan’ mammal, Lactodens sheni, from the Early Cretaceous Jehol Biota, represented by a partial skeleton with dentary and upper and lower teeth with dental morphologies well‐preserved. The new species has a dental formula of three upper incisors, one canine, three premolars, and six molars/three lower incisors, one canine, five premolars and six lower molars, double‐rooted canines, extremely low‐crowned and transversely thin premolars, and acute angled molars. The dental morphologies of molars and peculiar deciduous premolars are similar to those of Spalacolestes from North America. The associated upper and lower dentitions from one individual animal helped to clarify tooth identification of some spalacotheriids represented only by fragmentary material. Phylogenetic analyses indicate a close relationship of the new species to North American spalacolestines and faunal interchanges between Eurasia and North America, thus supporting the notion that small‐bodied spalacotheriids were diverse and had a pan‐Laurasian distribution during the Early Cretaceous. Absence of the Meckelian groove suggests acquisition of the definitive mammalian middle ear in spalacolestines, and deciduous canines and premolars in the slim and extremely long dentary imply a faunivorous diet.     

Patterns of size variation and correlation in the dentition of the red colobus monkey (Colobus badius)

There have been numerous studies on variability and correlation in dental crown size, but the significance of the resulting patterns remains unclear. Regions of low variation and high correlation have been hypothesized to represent the poles of Butler's morphological fields, to be related to absolute tooth size, or to be related to morphological complexity of the teeth and functional efficiency. Variation and correlation of tooth lengths and breadths were investigated in 138 red colobus monkeys to further assess the relations among size associations, crown morphology, and absolute tooth size. In the maxilla and mandible, the postcanine teeth are the most highly correlated and least variable, followed by the incisors, then the canines. There are also lower correlations between premolars and molars than within either group. While there appears to be a relation between degree of morphological differentiation and levels of correlation and variation, there are no notable differences in the correlation of opponents along the dental arcade, which is the most important functional consideration. This suggests that different levels of correlation and variation within upper or lower teeth are “artifacts” of tooth dimensions that contribute to different geometric designs in different tooth groups as the germs develop. This morphological effect is coupled with the influence of integration fields, indicated by higher variability and lower correlations of the third molar, the largest or most molarized tooth. It is concluded that there are wide functional tolerances in occlusion with respect to the gross dimensions of dental crowns and their interrelationships.     

Contours of maxillary molars studied in Australian aboriginals

Distances from the central pit to the perimeter of the crown of permanent upper molars were measured on standardized occlusal photographs of dental casts representing 210 male and 181 female Aboriginals from Yuendumu in the Northern Territory of Australia. In both males and females the first molar was the largest tooth but it showed least variability. Variabilities of the distances tended to be greater for radii constructed in the buccolingual direction than for the transverse mesiodistal radii. The most marked size reduction in the molar series from first to third related to the distolingual part of the crown, which was also the most variable region. Size differences between molars in the mesial contour radii were not marked. Sexual dimorphism was evident in most crown radii, being most marked for the second molar.     

Posterior dental size reduction in hominids: The Atapuerca evidence

In order to reassess previous hypotheses concerning dental size reduction of the posterior teeth during Pleistocene human evolution, current fossil dental evidence is examined. This evidence includes the large sample of hominid teeth found in recent excavations (1984–1993) in the Sima de los Huesos Middle Pleistocene cave site of the Sierra de Atapuerca (Burgos, Spain). The lower fourth premolars and molars of the Atapuerca hominids, probably older than 300 Kyr, have dimensions similar to those of modern humans. Further, these hominids share the derived state of other features of the posterior teeth with modern humans, such as a similar relative molar size and frequent absence of the hypoconulid, thus suggesting a possible case of parallelism. We believe that dietary changes allowed size reduction of the posterior teeth during the Middle Pleistocene, and the present evidence suggests that the selective pressures that operated on the size variability of these teeth were less restrictive than what is assumed by previous models of dental reduction. Thus, the causal relationship between tooth size decrease and changes in food-preparation techniques during the Pleistocene should be reconsidered. Moreover, the present evidence indicates that the differential reduction of the molars cannot be explained in terms of restriction of available growth space. The molar crown area measurements of a modern human sample were also investigated. The results of this study, as well as previous similar analyses, suggest that a decrease of the rate of cell proliferation, which affected the later-forming crown regions to a greater extent, may be the biological process responsible for the general and differential dental size reduction that occurred during human evolution. © 1995 Wiley-Liss, Inc.     

Family studies of tooth size factors in the permanent dentition.

Correlations between dimensions of the permanent teeth in Australian Aboriginals were studied by factor analysis to disclose the main sources of shared variability. Findings indicated that in both males and females most of the common variability in the tooth dimensions could be accounted for by factors representing mesiodistal size of anterior teeth, buccolingual size of anterior teeth, generalized size of the premolars and generalized size of the molars. Factor scores derived from the analysis were used to calculate intraclass correlations among brothers and among sisters. These correlations tended to be higher for the factors contributing most to the common variability indicating that the factors might represent fields under direct genetic control. There was no trend for intraclass correlations among siblings derived from multivariate scores to be consistently higher than those based on observed tooth dimensions. The main advantage to the user of factor analysis is the ability to interpret associations between interrelated variables more objectively than is possible by conventional correlation methods.     

Associations between Carabelli trait and cusp areas in human permanent maxillary first molars

Few dental anthropological studies have investigated the associations between tooth crown size and crown traits in humans using quantitative methods. We tested several hypotheses about overall crown size, individual cusp areas, and expression of Carabelli cusps in human permanent first molars by obtaining data from standardized occlusal photographs of 308 Australians of European descent (171 males and 137 females). Specifically, we aimed to calculate the areas of the four main molar cusps, and also Carabelli cusp, and to compare the relative variability of cusp areas in relation to timing of development. We also aimed to compare cusp areas between males and females and to describe how Carabelli cusp interacted with other molar cusps. Measurements included maximum crown diameters (mesiodistal and buccolingual crown diameters), the areas of the four main cusps, and the area of Carabelli cusp. The pattern of relative variability in absolute areas of molar cusps corresponded with their order of formation, the first-forming paracone displaying the least variation, and the last-forming Carabelli cusp showing the greatest. Overall crown size and areas of individual cusps all showed sexual dimorphism, with values in males exceeding those in females. Sexual dimorphism was smallest for paracone area and greatest for Carabelli cusp area. Overall crown size and cusp areas were larger in individuals displaying a Carabelli cusp, especially the hypocone area. Although the combined area of the protocone and a Carabelli cusp was greater in cuspal forms than noncuspal forms, protocone area alone was significantly smaller in the former. Our findings lead us to propose that, in individuals with the genotype for Carabelli trait expression, larger molar crowns are more likely to display Carabelli cusps, whereas molars with smaller crowns are more likely to display reduced forms of expression of the trait. We suggest that the pattern of folding of the internal enamel epithelium in developing molar crowns, particularly in the protocone region, can be modified by a developing Carabelli cusp.