Masticatory function and post-pleistocene evolution in Nubia

The present research focuses on craniofacial variation in Nubia over approximately 10,000 years. Samples were grouped according to their temporal location and subsistence pattern, and represent a transition from a hunting-gathering adaptation (Mesolithic) to a transitional hunting-gathering-agricultural adaptation (A-C Group) and finally to a fully agricultural adaptation (Meroitic/X-Group/Christian). The purposes were: (1) to compare the Mesolithic sample with the later Nubian populations; and (2) to evaluate further the hypothesis that change in Nubian craniofacial morphology was due to changing functional demands associated with the progressive change in subsistence adaptation and associated behavior. The results tend to support recent views that the Nubian Mesolithic population is probably ancestral to later Nubian groups, and that the masticatory-functional hypothesis can best account for craniofacial change among the Nubians since 12,000 B.P. According to this hypothesis systematic reduction in functional demand placed on the masticatory complex from the Mesolithic led, secondarily, to an alteration of the growth of the maxillomandibular complex such that the face became progressively less robust and more inferoposteriorly located relative to the cranial vault. Both the increase in the height of the vault relative to its length, producing a more “globular” appearance, and the reduction in dental size were tertiary, compensatory responses to altered facial size and position.     

Metric dental change in the European upper paleolithic and mesolithic.

Evolutionary trends for dental reduction are presented for European Upper Paleolithic and Mesolithic samples. The analysis demonstrates that the greatest decrease in tooth size occurs between the two divisions of the Upper Paleolithic, while little and insignificant change characterizes the Late Upper Paleolithic/Mesolithic transition. Trends for tooth size over this period indicate that (1) human evolution does not stop with the appearance of "anatomically modern Homo sapiens," (2) changes in tooth size fluctuate with increases in the efficiency and complexity of cultural systems, and (3) the Early Upper Paleolithic sample should be considered transitional between Wurm II European Neanderthals and later Upper Paleolithic and Mesolithic groups.     

Tooth Size Variation in Prehistoric India

Permanent dental crown dimensions are largely unknown for living and prehistoric populations of South Asia. This paper describes permanent tooth crown measurements and indices for the Late Chalcolithic skeletal series from Inamgaon (1700-700 B.C.) in western India. These data are compared with tooth size data for prehistoric populations in India and both living and prehistoric populations outside the South Asian subcontinent. In summed cross-sectional area, the Inamgaon (1,218 mm²) permanent teeth are most similar to prehistoric skeletal series from Mahurjhari, India, Non Nok Tha, Thailand, and Bellan Bandi Palassa, Sri Lanka. The Inamgaon total crown area is only 3.1% smaller than the figure reported for early Neolithic skeletons from Mehr-garh, Pakistan, indicating the relatively large size of the Inamgaon dentition. Large tooth size at Inamgaon is interpreted as a biological adaptation to coarse dietary items, basic food preparation methods, and a mixed economy that included hunting wild game and collecting wild fruit. Dental indices of "ethnic" significance are perhaps better interpreted as indicators of masticatory stress and differential dental reduction. Maxillary and mandibular third molars show minimal reduction in crown size, but mandibular second molar teeth are distinctly reduced in size. The Incisor Breadth Index may indicate mixed genetic ancestry of the Inamgaon people or similar dietary stresses in genetically different populations.     

A critique of the «increasing population density effect»

The «increasing population density effect» (IPDE) proposed by Macchiarelli and Bondioli (1986) represents one of the most recent attempts to account for dental reduction in modern human populations. Under the IPDE, the marked reduction in tooth size observed in post-Pleistocene human groups is seen as merely a side-effect of a more general reduction in body size, which resulted from an increase in population density. The model is dependent upon a strong correlation between tooth size and body size, which after numerous attempts has yet to be convincingly demonstrated in humans. This paper argues that the IPDE neglects the negative consequences on individual fitness of teeth which are too large to fit into diminishing jaws or are more susceptible to pathology, and that the worldwide reduction of tooth size is the result of selection for smaller teeth due to shifts to a softer and/or more cariogenic diet. Although increased population density may have intensified this selective effect by decreasing general fitness by lowering resistance to oral infections, it was not the primary cause of dental reduction. All proposed mechanisms of dental reduction, however, are in need of additional testing, and possible directions are offered for future studies of the phenomenon.     

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.     

Japanese tooth size: past and present

Mesial-distal and buccal-lingual crown measurements were made on male and female samples of recent Japanese teeth from three locations, Fukuoka, Kyoto, and Tokyo, and for Hokkaido Ainu and Koreans. Similar data were collected for prehistoric Middle-to-Late Jomon Japanese and from Yayoi specimens representing the first agriculturalists to appear in Japan. From a tooth-by-tooth comparison of cross-sectional areas, it was shown that the modern Japanese samples did not differ from one part of Japan to another. Korean tooth size also is not significantly different from Japanese, while Ainu have the smallest teeth recorded in Asia. The Yayoi who brought rice to Japan about 300 B.C. came in with teeth that were the same size as Chinese Neolithic teeth. They encountered a resident Jomon population whose teeth were 10% smaller. From tooth size measures alone, it is most economical to suggest that, if the rates of reduction observed elsewhere in the world applied in Japan, the recent Ainu would best be regarded as the direct descendants of the Jomon, while the modern Japanese are the results of in situ reduction from the incoming Yayoi. Other aspects of craniofacial morphology suggest that some Jomon was incorporated by the Yayoi. The modern Japanese, then, while predominantly derived from the Yayoi, would include a Jomon component.     

The effect of birthweight on tooth-size variability in twins.

Studies indicate that low birthweight (LBW) children display reduced deciduous tooth size but there is little information about permanent tooth size. It has also been shown that dental fluctuating asymmetry (FA) increases in response to various environmental influences, but the relationship between birthweight and FA remains unclear. The aim of this study was to compare tooth size and asymmetry, according to birthweight, in the deciduous and permanent dentitions of a sample of Australian twins. The study sample comprised 436 twins, classified into 2 groups: normal birthweight (NBW > 2500 g) and low birthweight (LBW < or = 2500 g). For each individual it was generally possible to measure maximum mesiodistal crown diameters of both deciduous and permanent central incisors from serial dental models. Correlations were calculated between tooth-size variables and birthweight; subsequently comparisons of tooth size and FA were made between the LBW and NBW samples using Student's t tests. Small positive correlations (around .1) were noted between birthweight and tooth-size variables. There was no evidence of tooth-size reduction in the LBW male sample, but the LBW females displayed tooth-size reduction of approximately 2-3% for both deciduous and permanent incisors, compared to the NBW females. There was no evidence of increased FA in the LBW individuals of either sex. These findings indicate that developing teeth are generally well-protected from developmental disturbances during prenatal and perinatal periods. Further research is needed to clarify the biological basis of an apparently true but weak association between tooth size and birthweight.     

Dental health of elderly confederate veterans: evidence from the Texas State Cemetery

Analysis of the skeletal remains of 50 Confederate veterans provided a unique opportunity to explore the dental health of a geriatric sample. These men, who died between 1907-1932, had an average age at death of 76.7 years. Ninety percent were institutionalized at the Confederate Home for Men (Austin, TX) prior to their deaths. This elderly sample was assessed in terms of caries, antemortem tooth loss (AMTL), abscesses, and linear enamel hypoplasias. On a per tooth basis, the AMTL rate was 57.2%. Of 39 dentate men, 33 (84.6%) had dental caries, and 24.4% (121 of 496) of teeth were carious. Ten (25.0%) of the dentate men had hypoplastic teeth. At least one abscess was seen in 14 (28%) of 50 individuals. Results from this geriatric institutionalized sample are compared to contemporaneous historical samples. Disparities in dental health among these groups may be due to differences in average age at death, and these comparisons allow a better understanding of dental changes that occur with age. The sample is also compared to modern elderly samples: modern groups have higher caries rates, possibly because they retained more teeth. This finding may be due in part to diets in the United States becoming increasingly cariogenic over time. In addition, dental care has moved from the reactive practices seen in the nineteenth and early twentieth centuries (such as tooth extractions) to modern proactive solutions dedicated to preserving and restoring teeth (such as tooth brushing, fluoride treatments, and dental fillings).     

Human dental reduction: natural selection or the probable mutation effect

Dental reduction has been sufficiently widespread among human populations to render the phenomenon of reduced tooth size worthy of scientific explanation. One of the most controversial models invoked to explain structural reduction in organisms is referred to as the "probable mutation effect" (PME). According to this model, structures no longer functional owing to ecological or cultural changes will experience a relaxation of selection pressure, permitting an accumulation of mutations in the population that inevitably will result in the reduction in size or the loss of the concerned structure. Although the PME continues to be offered as a viable explanation of human dental reduction, it is based upon several premises that modern dental clinical experience fails to support. Known enzyme defects resulting from mutations, factors predisposing to dental infections, and the deleterious effects of teeth that are too large or too small reveal that the PME does not logically account for the reduction of tooth size. Given such information, this paper proposes models of dental reduction based upon natural selection, which, unlike the PME, are testable in both modern and archaeological populations. The integration of clinical and skeletal data permits a more thorough understanding of dental reduction in the hominid fossil record.     

Post-pleistocene reductions in human dental structure: a reappraisal in terms of increasing population density

This paper discusses the well-documented acceleration which occurred in the reduction of human dental structure during the post-Pleistocene. It suggests that the process can be explained in terms of different but related factors inherent in a transition from late-Pleistocene hunting and gathering conditions to ones involving a sedentary life in larger groups. It is postulated that directional selective forces acting during Upper Paleolithic to maintain large tooth size had only a poor influence and that the new techniques in food preparation had little or no effect. The phenomenon is seen as a possible side-effect of a more complex overall reduction in body size, of which dental reduction only represents a small but demonstrable part. It is also suggested that stresses may have played an important role in producing a transitory reduced dimensional expression of the genetic background. An interaction between the variables directly or indirectly affecting body mass is shown in the model proposed which relates to post-Pleistocene. An increase in population density is indicated as being the most decisive biological factor in determining the acceleration in the trend towards reduction. A preliminary version of this paper was presented at the Symposium on Upper Paleolithic/Mesolithic of Europe and the Mediterranean Basin, Pisa, September 8–10, 1984.     

Human population differentiation in the peri-Baltic Mesolithic: the odontometrics of Oleneostrovskii mogilnik (Karelia)

Odontometric data are reported for Oleneostrovskii mogilnik, a Mesolithic cemetery in the Karelian USSR. Containing the skeletal remains of 148 individuals and dated to 7500b.p., it is the largest and oldest of the known peri-Baltic Mesolithic cemeteries. The dental data from this site are compared to those from Skateholm, where dental dimensions closely approximate the mean tooth sizes for all late Mesolithic European material. Olenii ostrov possesses a significantly smaller dentition, despite antedating the Skateholm sample by 1500 years. Geographic, archaeological, linguistic and ancillary dental data are discussed which argue strongly for the presence in Mesolithic northern Europe of at least two quasi-isolated human mating networks. One of these, documented by Olenii ostrov, may also represent the earliest evidence for the establishment in the taiga zone of populations whose descendants are the modern Saami (=”Lapps”).     

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.     

Population continuity vs. discontinuity revisited: dental affinities among late Paleolithic through Christian-era Nubians

The present study revisits a subject that has been a source of long-standing bioarchaeological contention, namely, estimation of Nubian population origins and affinities. Using the Arizona State University dental anthropology system, frequencies of 36 crown, root, and intraoral osseous discrete traits in 12 late Pleistocene through early historic Nubian samples were recorded and analyzed. Specifically, intersample phenetic affinities, and an indication of which traits are most important in driving this variation, were determined through the application of correspondence analysis and the mean measure of divergence distance statistic. The results support previous work by the author and others indicating that population discontinuity, in the form of replacement or significant gene flow into an existing gene pool, occurred sometime after the Pleistocene. This analysis now suggests that the break occurred before the Final Neolithic. Samples from the latter through Christian periods exhibit relative homogeneity, which implies overall post-Pleistocene diachronic and regional population continuity. Yet there are several perceptible trends among these latter samples that: 1) are consistent with documented Nubian population history, 2) enable the testing of several existing peopling hypotheses, and 3) allow the formulation of new hypotheses, including a suggestion of two post-Pleistocene subgroups predicated on an age-based sample dichotomy.     

Increase of tooth size in prehistoric coastal Peru, 10,000 B.P.-1,000 B.P.

Teeth increase in size during a 9,000-year period in an archaeologically derived, radiocarbon dated sample of skeletons from a geographically restricted area of coastal Peru. Although cultural change is extensive, including the transition to food production and pottery making, teeth do not reduce as predicted under these conditions by Brace's Probable Mutation Effect. Since most of the dental literature dealing with size change of teeth focuses upon dental reduction, hypotheses explaining why teeth increase through time are not well developed. No obvious selective forces explaining size increase are apparent in the present data. Attrition decreases through time. The increase in tooth size in this collection may be a function of overall cranialfacial size increase, which (pending further data) may be related to a general body size increase.     

Patterning the size and number of tooth and its cusps

Mice and rats, two species of rodents, show some dental similarities such as tooth number and cusp number, and differences such as tooth size and cusp size. In this study, the tooth size, tooth number, cusp size and cusp number, which are four major factors of the tooth patterning, were investigated by the heterospecific recombinations of tissues from the molar tooth germs of mice and rats. Our results suggest that the dental epithelium and mesenchyme determine the cusp size and tooth size respectively and the cusp number is co-regulated by the tooth size and cusp size. It is also suggested that the mesenchymal cell number regulates not the tooth size but the tooth number. The relationships among these factors in tooth patterning including micropatterning (cusp size and cusp number) and macropatterning (tooth size and tooth number) were analyzed in a reaction diffusion mechanism. Key molecules determining the patterning of teeth remains to be elucidated for controlling the tooth size and cusp size of bioengineered tooth.     

Post-Pleistocene facial reduction,biomechanics and selection against morphologically complex teeth: A rejoinder to macchiarelli and bondioli

Macchiarelli & Bondioli (1984, 1986) argue that post-Pleistocene cranio-facial reduction cannot be explained by biomechanical factors related to change in the diet (Carlson, 1974;Carlson & Van Gerven, 1977) or to facial reduction related to the selective advantages of smaller, morphologically simpler teeth (Greene, 1967;Van Gerven, Armelagos & Rohr, 1977). Instead, they maintain that facial reduction is a mere side-effect of a reduction in overall body size. Our analysis of skeletal and facial reduction in post-Pleistocene Sudanese Nubia suggests thatMacchiarelli & Bondioli's interpretation is incorrect for two reasons. First, the reduction in facial morphology (at least since Mesolithic times) is greater by several orders of magnitude than the reduction in general body size. Second, the dentition not only shows a greater size reduction than does general body size, but a shifting pattern of dental reduction rather than a general decrease across all teeth.     

The dentition of the Indian Knoll skeletal population: odontometrics and cusp number.

Data on the permanent dentition of 153 individuals from the well known Indian Knoll skeletal population are presented. Mesiodistal and buccolingual measurements were taken with a Helios dial caliper. Cusp number of maxillary and mandibular molars are recorded. The Indian Knoll dentition is larger than many modern groups but smaller than Australoid or Mesolithic groups. With the exception of maxillary 12, males have larger teeth than females in both dimensions. The lower canine is the most dimorphic tooth. Through rank order correlation, an association was shown between the sexual dimorphism of the mesiodistal and buccolingual dimensions. Compared to modern groups, the Indian Knoll population displays a moderate degree of sexual dimorphism in tooth size. In general, the coefficients of variation were greater for the more distal teeth within morphological classes. Amounts of size variability did not differ significantly between the sexes; moreover, rank order correlations indicated that patterns of variability in both dimensions were similar for males and females. The predominant cusp number pattern for upper molars is 4-3-3 and for lowers 5-5(4)-5. No sex differences were shown for cusp occurrence or bilateral asymmetry in cusp number.     

Temporal variation in prehistoric Nubian crania

Much of the earlier work on the prehistory of Sudanese Nubia has emphasized discontinuity between early Nubian populations. However, recent investigations suggest the converse - that a remarkable degree of cultural and biological continuity exists among indigenous Nubian groups, perhaps as far back as the Paleolithic. Thus, cultural and biological differences between Nubian populations can be most effectively perceived as the result of in situ evolutionary development. The present analysis has two major purposes: (1) to describe the morphological differences in the craniofacial complex between indigenous Nubian populations extending from the A-Group (c. 3,400 B.C.) through the Christian (c. 1,500 A.D.) horizons; and (2) to account for these differences within an evolutionary framework. The multiple discriminant analysis of radiographically derived variables revealed a trend from a substantially lower and more elongated cranial vault to a shorter and taller vault throughout the almost 5,000 year time span. Associated with this pattern was a tendency for the face to become more inferiorly-posteriorly located with respect to the vault in the latter groups. Finally, the masseter and temporalis muscles underwent a reduction and slight relocation through time. We speculate that this trend may be associated with behavioral changes associated with transition from a hunting and gathering to a totally agricultural subsistence pattern.     

Crown dimensions of deciduous teeth from prehistoric India

Mean mesiodistal and buccolingual diameters are presented for deciduous teeth from the important Chalcolithic site of Inamgaon (1400–700 B.C.), a prehistoric farming community on the Deccan Plateau of western India. The deciduous teeth from Inamgaon are consistently larger than deciduous teeth of modern populations of European descent and smaller than the deciduous teeth of modern Australian aboriginals. Comparative data for prehistoric deciduous teeth are rare, especially for populations of southern Asia. The deciduous teeth of Mesolithic Europeans are comparable in size to certain dimensions of the Inamgaon teeth, and a small sample of deciduous teeth from the Iron Age site of Pomparippu (Sri Lanka) exhibits larger anterior teeth and smaller molar teeth than does the sample from Inamgaon.