A functional analysis of Carabelli trait in Australian aboriginal dentition.

OBJECTIVES: Carabelli is a nonmetric dental trait variably expressed as a small pit to a prominent cusp in the maxillary molars of modern humans. Investigations on the occurrence and expression rates of this trait have been conducted extensively, tracing its origin to genetic sources. However, there remains a lack of understanding about its potential role in chewing. In this study, we examine molar macrowear with the aim of reconstructing Carabelli trait occlusal dynamics occurring during chewing. METHODS: We have examined 96 deciduous and permanent maxillary molars of children and young adults from Yuendumu, an Australian Aboriginal population that was at an early stage of transition from a nomadic and hunter-gatherer way of life to a more settled existence. We apply a well-established method, called Occlusal Fingerprint Analysis, which is a digital approach for analyzing dental macrowear allowing the reconstruction of jaw movements required to produce wear pattern specific to each tooth. RESULTS: Carabelli trait slightly enlarges the surface functional area, especially in those molars where this feature is expressed in its cuspal form and it is closer to the occlusal plane. Moreover, the highly steep contact planes would also indicate that Carabelli wear areas contribute to increasing the shearing abilities of the occluded teeth, which are particularly important when processing fibrous and tough foods. CONCLUSIONS: The macrowear analysis suggests that Carabelli trait in the Aboriginal people from Yuendumu slightly enhanced occlusion and probably played some functional role during mastication. Future biomechanical and microwear analyses could provide additional information on the mechanical adaptation of Carabelli trait in modern human dentition.

A mathematical landmark-based method for measuring worn molars in hominoid systematics.

Worn teeth pose a major limitation to researchers in the fields of extinct and extant hominoid systematics because they lack clearly identifiable anatomical landmarks needed to take measurements on the crown enamel surface and are typically discarded from a study. This is particularly detrimental when sample sizes for some groups are already characteristically low, if there is an imbalance between samples representing populations, sexes or dietary strategies, or if the worn teeth in question are type specimens of fossil species or other key specimens. This study proposes a methodology based predominantly on mathematically-derived landmarks for measuring size and shape features of molars, irrespective of wear. With 110 specimens of lower second molars from five species of extant hominoids (Pan troglodytes, P. paniscus, Gorilla gorilla, G. beringei, Homo sapiens), n ≥ 20 per species, n ≥ 10 per subspecies, good species separation in morphospace is achieved in a principal components analysis. Classification accuracy in a discriminant function analysis is 96.4% at the species level and 88.2% at the subspecies level (92.7% and 79.1%, respectively, on cross-validation). The classification accuracy compares favorably to that achieved by anatomically-derived measurements based on published research (94% and 84% at the species and subspecies level respectively; 91% and 76% on cross-validation). The mathematical landmarking methodology is rapid and uncomplicated. The results support the use of mathematical landmarks to enable the inclusion of worn molar teeth in dental studies so as to maximize sample sizes and restore balance between populations and/or sexes in hominoid systematic studies.

Generalized Procrustes analysis of an ontogenetic series of modified crania: Evaluating the technique of modification in the Migration Period of Europe (4th-7th century AD).

OBJECTIVES: The arrival of the Huns into Europe in the fourth century AD increased the occurrence of intentional cranial modification among European nomads. It has been postulated that the Huns used a two-bandage cranial binding technique to differentiate themselves from surrounding nomadic groups, including those from Georgia. This study examines this hypothesis by comparing Migration Period (4th to 7th century AD) juvenile crania, which retain strong impressions of bindings, with adult modified crania from Hungary and Georgia. MATERIALS AND METHODS: Twelve surface landmarks and 251 semi-landmarks were used to study ontogenetic trajectories in 9 juvenile and 16 adult modified skulls from 8 Hungarian sites and 21 adult skulls from two Georgian sites. Generalized Procrustes analysis, linear regression of Procrutes distance on dental age and log centroid size, and warping the principal components (PCs) in shape space helped to identify cranial shape changes. RESULTS: The PCs provide significant separation of the juvenile and adult groups from Georgia and Hungary. Variation in modified cranial shape was limited in Hungary compared to Georgia. There was stronger correlation between juvenile and adult modified cranial shape in Hungary than in Georgia. Warping along the first axis reveals the trajectory from marked flattening of the frontal and occipital regions in juveniles to diminished flattening in the same regions in adult crania, corresponding with one binding. Another depression extending from the post-bregmatic region to the temporal region, similarly strong in juveniles but diminishing in adults, marks the second binding. DISCUSSION: Hungarian crania were modified with two bindings with limited shape variation, whereas the Georgian crania had greater variation in shape being also modified with antero-posterior bindings. The findings from this study alongside contemporary historical sources help to understand the role of intentional cranial modification as a mark of social identity among nomads in the Migration Period of Europe.

Migrating Huns and modified heads: Eigenshape analysis comparing intentionally modified crania from Hungary and Georgia in the Migration Period of Europe.

An intentionally modified head is a visually distinctive sign of group identity. In the Migration Period of Europe (4th- 7th century AD) the practice of intentional cranial modification was common among several nomadic groups, but was strongly associated with the Huns from the Carpathian Basin in Hungary, where modified crania are abundant in archaeological sites. The frequency of modified crania increased substantially in the Mtskheta region of Georgia in this time period, but there are no records that Huns settled here. We compare the Migration Period modified skulls from Georgia with those from Hungary to test the hypothesis that the Huns were responsible for cranial modification in Georgia. We use extended eigenshape analysis to quantify cranial outlines, enabling a discriminant analysis to assess group separation and identify morphological differences. Twenty-one intentionally modified skulls from Georgia are compared with sixteen from Hungary, using nineteen unmodified crania from a modern population as a comparative baseline. Results indicate that modified crania can be differentiated from modern unmodified crania with 100% accuracy. The Hungarian and Georgian crania show some overlap in shape, but can be classified with 81% accuracy. Shape gradations along the main eigenvectors indicate that the Hungarian crania show little variation in cranial shape, in accordance with a two-bandage binding technique, whereas the Georgian crania had a wider range of variation, fitting with a diversity of binding styles. As modification style is a strong signifier of social identity, our results indicate weak Hunnic influence on cranial modification in Georgia and are equivocal about the presence of Huns in Georgia. We suggest instead that other nomadic groups such as Alans and Sarmatians living in this region were responsible for modified crania in Georgia.

The Taxonomic and Phylogenetic Affinities of Bunopithecus sericus, a Fossil Hylobatid from the Pleistocene of China.

Fossil hylobatids are rare, but are known from late Miocene and Pleistocene sites throughout East Asia. The best-known fossil hylobatid from the Pleistocene of China is a left mandibular fragment with M2-3 (AMNH 18534), recovered from a pit deposit near the village of Yanjinggou in Wanzhou District, Chongqing Province. Matthew and Granger described this specimen in 1923 as a new genus and species, Bunopithecus sericus. Establishing the age of Bunopithecus has proved difficult because the Yanjinggou collection represents a mixed fauna of different ages, but it likely comes from early or middle Pleistocene deposits. Although the Bunopithecus specimen has featured prominently in discussions of hylobatid evolution and nomenclature, its systematic status has never been satisfactorily resolved. The present study reexamines the taxonomic and phylogenetic relationships of Bunopithecus by carrying out a detailed comparative morphometric study of its lower molars in relation to a large sample of modern hylobatids. Our results show that differences in M2 and M3 discriminate extant hylobatids fairly well, at least at the generic level, and that AMNH 18534 is not attributable to Hylobates, Nomascus or Symphalangus. Support for a close relationship between Bunopithecus and Hoolock is more equivocal. In most multivariate analyses, Bunopithecus presents a unique morphological pattern that falls outside the range of variation of any hylobatid taxon, although its distance from the cluster represented by extant hoolocks is relatively small. Our results support the generic distinction of Bunopithecus, which most likely represents an extinct crown hylobatid, and one that may possibly represent the sister taxon to Hoolock.

Using CRANID to test the population affinity of known crania.

CRANID is a statistical program used to infer the source population of a cranium of unknown origin by comparing its cranial dimensions with a worldwide craniometric database. It has great potential for estimating ancestry in archaeological, forensic and repatriation cases. In this paper we test the validity of CRANID in classifying crania of known geographic origin. Twenty-three crania of known geographic origin but unknown sex were selected from the osteological collections of the University of Melbourne. Only 18 crania showed good statistical match with the CRANID database. Without considering accuracy of sex allocation, 11 crania were accurately classified into major geographic regions and nine were correctly classified to geographically closest available reference populations. Four of the five crania with poor statistical match were nonetheless correctly allocated to major geographical regions, although none was accurately assigned to geographically closest reference samples. We conclude that if sex allocations are overlooked, CRANID can accurately assign 39% of specimens to geographically closest matching reference samples and 48% to major geographic regions. Better source population representation may improve goodness of fit, but known sex-differentiated samples are needed to further test the utility of CRANID.

Dental and phylogeographic patterns of variation in gorillas.

Gorilla patterns of variation have great relevance for studies of human evolution. In this study, molar morphometrics were used to evaluate patterns of geographic variation in gorillas. Dental specimens of 323 adult individuals, drawn from the current distribution of gorillas in equatorial Africa were divided into 14 populations. Discriminant analyses and Mahalanobis distances were used to study population structure. Results reveal that: 1) the West and East African gorillas form distinct clusters, 2) the Cross River gorillas are well separated from the rest of the western populations, 3) gorillas from the Virunga mountains and the Bwindi Forest can be differentiated from the lowland gorillas of Utu and Mwenga-Fizi, 4) the Tshiaberimu gorillas are distinct from other eastern gorillas, and the Kahuzi-Biega gorillas are affiliated with them. These findings provide support for a species distinction between Gorilla gorilla and Gorilla beringei, with subspecies G. g. diehli, G. g. gorilla, G. b. graueri, G. b. beringei, and possibly, G. b. rex-pygmaeorum. Clear correspondence between dental and other patterns of taxonomic diversity demonstrates that dental data reveal underlying genetic patterns of differentiation. Dental distances increased predictably with altitude but not with geographic distances, indicating that altitudinal segregation explains gorilla patterns of population divergence better than isolation-by-distance. The phylogeographic pattern of gorilla dental metric variation supports the idea that Plio-Pleistocene climatic fluctuations and local mountain building activity in Africa affected gorilla phylogeography. I propose that West Africa comprised the historic center of gorilla distribution and experienced drift-gene flow equilibrium, whereas Nigeria and East Africa were at the periphery, where climatic instability and altitudinal variation promoted drift and genetic differentiation. This understanding of gorilla population structure has implications for gorilla conservation, and for understanding the distribution of sympatric chimpanzees and Plio-Pleistocene hominins.

Population systematics of chimpanzees using molar morphometrics.

When dental morphological variation within extant species is used as a guideline to partition variation within fossil samples into species, the underlying assumption is that fossil species are equivalent to extant species. This is the case despite the fact that dental morphology, which is commonly used to differentiate fossil species, is rarely used to differentiate extant species. Aspects of external morphology, ecology, behavior, breeding patterns, and molecular structure that are used to delineate living species are generally not available for fossils. In this paper, the utility of dental evidence for sorting fossil samples into species is evaluated by testing whether molar occlusal morphology is capable of sorting populations of Pan into the species and subspecies already well-established by nondental evidence. The dentitions of 341 chimpanzee individuals, sampled from regions throughout equatorial Africa, were sorted into 16 populations using rivers to demarcate the boundaries between populations. Digital-imaging software was used to measure 15 traits on the occlusal surface of each upper molar and 19 on each lower molar. After applying size adjustments, size-transformed and untransformed variables were subjected to discriminant analysis, with separate analyses carried out for each molar type. Results indicate that populations of Pan troglodytes and Pan paniscus are well differentiated at all molar positions. Populations of P. t. verus are distinct from other populations of P. troglodytes. Populations of P. t. troglodytes and P. t. schweinfurthii show close dental similarity. A distinct population is recognized at the Nigeria-Cameroon border, indicating the presence of P. t. vellerosus. The concordance between the patterns of diversity recognized by this study and other molecular and nonmolecular studies indicates that paleontological species that are similar to species of Pan in terms of size and patterns of diversification may be differentiated using molar morphology.

Lingual incisor traits in modern hominoids and an assessment of their utility for fossil hominoid taxonomy.

The morphology of the anterior dentition has received scant attention for purposes of taxonomic discrimination. Recently, however, lingual incisor morphology was used in differentiating several Miocene ape species and genera. This paper assesses the utility of this morphology for taxonomic discrimination by examining the nature and patterns of variation in lingual incisor morphology in extensive samples of modern chimpanzees, gorillas, orangutans, and gibbons. This paper documents discrete morphological traits on the lingual side of incisors. Trait frequencies are used in univariate and multivariate analyses to examine the apportionment of variation in species, subspecies, and populations. A correlation between lingual incisor traits, tooth dimensions, and sex attempts to determine if such factors affect the manifestation of traits. Finally, the findings are applied to understanding patterns of variation in the Miocene hominids. The study demonstrates that: 1) lingual incisor morphology differs substantially between the hylobatids and great apes; 2) variation in incisor traits is high within species, and most of it is found within local populations; and 3) incisor traits do not correlate significantly with incisor dimensions or sex. Species and to some extent subspecies of extant hominoids can be differentiated statistically using lingual incisor traits, but the frequency of traits such as continuous or discontinuous cingulum, or the presence or absence of pillars, differentiates them. Given this pattern of variation, I argue that it is necessary to assume and document similar patterns of variation in Miocene apes before incisor morphology is used for differentiating taxa.

Interobserver error involved in independent attempts to measure cusp base areas of Pan M1s.

Cusp base areas measured from digitized images increase the amount of detailed quantitative information one can collect from post-canine crown morphology. Although this method is gaining wide usage for taxonomic analyses of extant and extinct hominoids, the techniques for digitizing images and taking measurements differ between researchers. The aim of this study was to investigate interobserver error in order to help assess the reliability of cusp base area measurement within extant and extinct hominoid taxa. Two of the authors measured individual cusp base areas and total cusp base area of 23 maxillary first molars (M(1)) of Pan. From these, relative cusp base areas were calculated. No statistically significant interobserver differences were found for either absolute or relative cusp base areas. On average the hypocone and paracone showed the least interobserver error (< 1%) whereas the protocone and metacone showed the most (2.6-4.5%). We suggest that the larger measurement error in the metacone/protocone is due primarily to either weakly defined fissure patterns and/or the presence of accessory occlusal features. Overall, levels of interobserver error are similar to those found for intraobserver error. The results of our study suggest that if certain prescribed standards are employed then cusp and crown base areas measured by different individuals can be pooled into a single database.