Ottmar Kullmer

Early dispersal of modern humans in Europe and implications for Neanderthal behaviour

The appearance of anatomically modern humans in Europe and the nature of the transition from the Middle to Upper Palaeolithic are matters of intense debate. Most researchers accept that before the arrival of anatomically modern humans, Neanderthals had adopted several ‘transitional’ technocomplexes. Two of these, the Uluzzian of southern Europe and the Châtelperronian of western Europe, are key to current interpretations regarding the timing of arrival of anatomically modern humans in the region and their potential interaction with Neanderthal populations. They are also central to current debates regarding the cognitive abilities of Neanderthals and the reasons behind their extinction. However, the actual fossil evidence associated with these assemblages is scant and fragmentary, and recent work has questioned the attribution of the Châtelperronian to Neanderthals on the basis of taphonomic mixing and lithic analysis. Here we reanalyse the deciduous molars from the Grotta del Cavallo (southern Italy), associated with the Uluzzian and originally classified as Neanderthal. Using two independent morphometric methods based on microtomographic data, we show that the Cavallo specimens can be attributed to anatomically modern humans. The secure context of the teeth provides crucial evidence that the makers of the Uluzzian technocomplex were therefore not Neanderthals. In addition, new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to ∼45,000–43,000 calendar years before present. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals.

Using occlusal wear information and finite element analysis to investigate stress distributions in human molars

Simulations based on finite element analysis (FEA) have attracted increasing interest in dentistry and dental anthropology for evaluating the stress and strain distribution in teeth under occlusal loading conditions. Nonetheless, FEA is usually applied without considering changes in contacts between antagonistic teeth during the occlusal power stroke. In this contribution we show how occlusal information can be used to investigate the stress distribution with 3D FEA in lower first molars (M1). The antagonistic crowns M1 and P2–M1 of two dried modern human skulls were scanned by μCT in maximum intercuspation (centric occlusion) contact. A virtual analysis of the occlusal power stroke between M1 and P2–M1 was carried out in the Occlusal Fingerprint Analyser (OFA) software, and the occlusal trajectory path was recorded, while contact areas per time‐step were visualized and quantified. Stress distribution of the M1 in selected occlusal stages were analyzed in strand7, considering occlusal information taken from OFA results for individual loading direction and loading area. Our FEA results show that the stress pattern changes considerably during the power stroke, suggesting that wear facets have a crucial influence on the distribution of stress on the whole tooth. Grooves and fissures on the occlusal surface are seen as critical locations, as tensile stresses are concentrated at these features. Properly accounting for the power stroke kinematics of occluding teeth results in quite different results (less tensile stresses in the crown) than usual loading scenarios based on parallel forces to the long axis of the tooth. This leads to the conclusion that functional studies considering kinematics of teeth are important to understand biomechanics and interpret morphological adaptation of teeth.

Molar Macrowear Reveals Neanderthal Eco-Geographic Dietary Variation

Neanderthal diets are reported to be based mainly on the consumption of large and medium sized herbivores, while the exploitation of other food types including plants has also been demonstrated. Though some studies conclude that early Homo sapiens were active hunters, the analyses of faunal assemblages, stone tool technologies and stable isotopic studies indicate that they exploited broader dietary resources than Neanderthals. Whereas previous studies assume taxon-specific dietary specializations, we suggest here that the diet of both Neanderthals and early Homo sapiens is determined by ecological conditions. We analyzed molar wear patterns using occlusal fingerprint analysis derived from optical 3D topometry. Molar macrowear accumulates during the lifespan of an individual and thus reflects diet over long periods. Neanderthal and early Homo sapiens maxillary molar macrowear indicates strong eco-geographic dietary variation independent of taxonomic affinities. Based on comparisons with modern hunter-gatherer populations with known diets, Neanderthals as well as early Homo sapiens show high dietary variability in Mediterranean evergreen habitats but a more restricted diet in upper latitude steppe/coniferous forest environments, suggesting a significant consumption of high protein meat resources.

Technical note: Occlusal fingerprint analysis: Quantification of tooth wear pattern

Information about food ingestion and mastication behavior during the lifespan of an individual is encoded in the dental occlusal wear pattern. To decode this information, we describe a new method called occlusal fingerprint analysis (OFA). Structural parameters of wear facets on the occlusal surface of teeth are quantified from digitized casts for the interpretation of occlusal aspects. The OFA provides an individual three-dimensional dental occlusal compass that indicates the major pathways of interaction between antagonists, revealing information about development, spatial position, and enlargement of wear facets. Humans develop a very similar overall pattern of crown contacts, although specific characteristics of wear facets reflect an individuals occlusal relationships and masticatory behavior. We hypothesize that the wear pattern is a unique character and therefore valuable for individual identification. Furthermore we suggest that OFA, when further developed, may be useful for identification of behavioral, biological, and chemical factors affecting crown morphology.

Comparison of dental measurement systems for taxonomic assignment of first molars

Morphometrics of the molar crown is based traditionally on diameter measurements but is nowadays more often based on 2D image analysis of crown outlines. An alternative approach involves measurements at the level of the cervical line. We compare the information content of the two options in a three-dimensional (3D) digital sample of lower and upper first molars (M(1) and M(1) ) of modern human and Neanderthal teeth. The cervical outline for each tooth was created by digitizing the cervical line and then sectioning the tooth with a best fit plane. The crown outline was projected onto this same plane. The curves were analyzed by direct extraction of diameters, diagonals, and area and also by principal component analysis either of the residuals obtained by regressing out these measurements from the radii (shape information) or directly by the radii (size and shape information). For M(1) , the crown and cervical outline radii allow us to discriminate between Neanderthals and modern humans with 90% and 95% accuracy, respectively. Fairly good discrimination between the groups (80-82.5%) was also obtained using cervical measurements. With respect to M(1) , general overlap of the two groups was obtained by both crown and cervical measurements; however, the two taxa were differentiable by crown outline residuals (90-97%). Accordingly, while crown diameters or crown radii should be used for taxonomic analysis of unworn or slightly worn M(1) s, the crown outline, after regressing out size information, could be promising for taxonomic assignment of lower M1s.

A new 3-d approach to determine functional morphology of cercopithecoid molars

Functional relationships between diet and tooth morphology form an integral part of primatological and paleontological research. Previously, mostly two-dimensional parameters have been used to compare and interpret the complex crown morphology of cercopithecine and colobine molars. However, as teeth are three-dimensional objects, any dimensional reduction in describing their morphology must result in loss of information. In the current study we use a high resolution optical topometric system to record crown morphology in different wear stages in order to extract three-dimensional (3-d) parameters from virtual 3-d models. Structural parameters such as relief index, occlusal surface area, enamel area and strike and dip of cusp slopes as well as wear facets can be calculated, reflecting the changing occlusal topography of molars due to attrition and abrasion. By comparing mostly fruit-eating cercopithecines and leaf-eating colobines, functional implications of tooth wear, occlusal jaw movement and resulting relief can be inferred. Our analyses show that the main differences in occlusal relief between these genera do not change with wear. Colobines maintain higher occlusal relief, whereas in cercopithecines all cusps wear flat quite rapidly. Detailed models of wear for cercopithecoid molars will be used to reconstruct diet and will enhance our knowledge of the paleoecology of Cercopithecoidea.

Comparison of occlusal loading conditions in a lower second premolar using three-dimensional finite element analysis.

ObjectivesThis study aimed to compare the patterns of stress distribution in a lower second premolar using three conventional occlusal loadings and two more realistic loading scenarios based on occlusal contact areas.Materials and methodsThe teeth of a dried modern human skull were micro-CT scanned in maximum intercuspation contact with a Viscom X8060 NDT X-ray system. A kinematic analysis of the surface contacts between antagonistic right upper and lower teeth during the power stroke was carried out in the Occlusal Fingerprint Analyser (OFA) software. Stress distribution in the lower right second premolar was analysed using three-dimensional finite element (FE) methods, considering occlusal information taken from OFA results (cases 4–5). The output was compared to that obtained by loading the tooth with a single point force (cases 1–3).ResultsResults for cases 1–3 differ considerable from those of cases 4–5. The latter show that tensile stresses might be concentrated in grooves and fissures of the occlusal surface, in the marginal ridges, in the disto-lingual and in the distal side of the root. Moreover, the premolar experiences high tensile stresses in the buccal aspect of the crown, supporting the idea that abfraction might be a dominant factor in the aetiology of non-carious cervical lesions.ConclusionsThe application of FE methods in dental biomechanics can be advanced considering individual wear patterns.Clinical relevanceMore realistic occlusal loadings are of importance for both new developments in prosthetic dentistry and improvements of materials for tooth restoration, as well to address open questions about the worldwide spread problem of dental failure.

Brief communication: Comparing loading scenarios in lower first molar supporting bone structure using 3D finite element analysis

Finite element analysis (FEA) is a widespread technique to evaluate the stress/strain distributions in teeth or dental supporting tissues. However, in most studies occlusal forces are usually simplified using a single vector (i.e., point load) either parallel to the long tooth axis or oblique to this axis. In this pilot study we show how lower first molar occlusal information can be used to investigate the stress distribution with 3D FEA in the supporting bone structure. The LM(1) and the LP(2) -LM(1) of a dried modern human skull were scanned by μCT in maximum intercuspation contact. A kinematic analysis of the surface contacts between LM(1) and LP(2) -LM(1) during the power stroke was carried out in the occlusal fingerprint analyzer (OFA) software to visualize contact areas during maximum intercuspation contact. This information was used for setting the occlusal molar loading to evaluate the stress distribution in the supporting bone structure using FEA. The output was compared to that obtained when a point force parallel to the long axis of the tooth was loaded in the occlusal basin. For the point load case, our results indicate that the buccal and lingual cortical plates do not experience notable stresses. However, when the occlusal contact areas are considered, the disto-lingual superior third of the mandible experiences high tensile stresses, while the medio-lingual cortical bone is subjected to high compressive stresses. Developing a more realistic loading scenario leads to better models to understand the relationship between masticatory function and mandibular shape and structures. Am J Phys Anthropol, 2012.

To meat or not to meat? New perspectives on Neanderthal ecology.

Neanderthals have been commonly depicted as top predators who met their nutritional needs by focusing entirely on meat. This information mostly derives from faunal assemblage analyses and stable isotope studies: methods that tend to underestimate plant consumption and overestimate the intake of animal proteins. Several studies in fact demonstrate that there is a physiological limit to the amount of animal proteins that can be consumed: exceeding these values causes protein toxicity that can be particularly dangerous to pregnant women and newborns. Consequently, to avoid food poisoning from meat-based diets, Neanderthals must have incorporated alternative food sources in their daily diets, including plant materials as well.

Comparison of dental measurement systems for taxonomic assignment of Neanderthal and modern human lower second deciduous molars

Traditional morphometric approaches for taxonomic assignment of Neanderthal and modern human dental remains are mainly characterized by caliper measurements of tooth crowns. Several studies have recently described differences in dental tissue proportions and enamel thickness between Neanderthal and modern human teeth. At least for the lower second deciduous molar (dm(2)), a three-dimensional lateral relative enamel thickness index has been proposed for separating the two taxa. This index has the advantage over other measurements of being applicable to worn teeth because it ignores the occlusal aspect of the crown. Nevertheless, a comparative evaluation of traditional crown dimensions and lateral dental tissue proportion measurements for taxonomic assignment of Neanderthal and modern human dm(2)s has not yet been performed. In this study, we compare various parameters gathered from the lateral aspects of the crown. These parameters include crown diameters, height of the lateral wall of the crown (lateral crown height = LCH), lateral enamel thickness, and dentine volume of the lateral wall, including the volume of the coronal pulp chamber (lateral dentine plus pulp volume = LDPV), in a 3D digital sample of Neanderthal and modern human dm(2)s to evaluate their utility in separating the two taxa. The LDPV and the LCH allow us to discriminate between Neanderthals and modern humans with 88.5% and 92.3% accuracy, respectively. Though our results confirm that Neanderthal dm(2)s have lower relative enamel thickness (RET) index compared with modern humans (p = 0.005), only 70% of the specimens were correctly classified on the basis of the RET index. We also emphasize that results of the lateral enamel thickness method depend on the magnitude of the interproximal wear. Accordingly, we suggest using the LCH or the LDPV to discriminate between Neanderthal and modern human dm(2)s. These parameters are more independent of interproximal wear and loss of lateral enamel.