New permanent teeth from Gran Dolina-TD6 (Sierra de Atapuerca). The bearing of Homo antecessor on the evolutionary scenario of Early and Middle Pleistocene Europe

Here we analyze the unpublished hominin dental remains recovered from the late Early Pleistocene Gran Dolina-TD6.2 level of the Sierra de Atapuerca (northern Spain), as well as provide a reassessment of the whole TD6.2 hominin dental sample. Comparative descriptions of the outer enamel surface (OES) and the enamel-dentine junction (EDJ) are provided. Overall, the data presented here support the taxonomic validity of Homo antecessor, since this species presents a unique mosaic of traits. Homo antecessor displays several primitive features for the genus Homo as well as some traits exclusively shared with Early and Middle Pleistocene Eurasian hominins. Some of these Eurasian traits were retained by the Middle Pleistocene hominins of Europe, and subsequently became the typical condition of the Neanderthal lineage. Although other skeletal parts present resemblances with Homo sapiens, TD6.2 teeth do not show any synapomorphy with modern humans. In addition, TD6.2 teeth can be well differentiated from those of Asian Homo erectus. The dental evidence is compatible with previous hypothesis about H. antecessor belonging to the basal population from which H. sapiens, Homo neanderthalensis, and Denisovans emerged. Future findings and additional research may help to elucidate the precise phylogenetic link among them.

First systematic assessment of dental growth and development in an archaic hominin (genus, Homo) from East Asia

Several human dental traits typical of modern humans appear to be associated with the prolonged period of development that is a key human attribute. Understanding when, and in which early hominins, these dental traits first appeared is thus of strong interest. Using x-ray multiresolution synchrotron phase-contrast microtomography, we quantify dental growth and development in an archaic Homo juvenile from the Xujiayao site in northern China dating to 161,000–224,000 years or 104,000–125,000 years before present. Despite the archaic morphology of Xujiayao hominins, most aspects of dental development of this juvenile fall within modern human ranges (e.g., prolonged crown formation time and delayed first molar eruption). For its estimated age-at-death (6.5 years), its state of dental development is comparable to that of equivalently aged modern children. These findings suggest that several facets of modern human dental growth and development evolved in East Asia before the appearance of fully modern human morphology. An archaic Homo juvenile from the East Asian Middle-Late Pleistocene transition has surprisingly modern dental development. An archaic Homo juvenile from the East Asian Middle-Late Pleistocene transition has surprisingly modern dental development.

Enamel and dentine dimensions of the Pleistocene hominins from Atapuerca (Burgos, Spain): A comparative study of canine teeth

Enamel and dentin patterns have awakened a considerable interest in phylogenetic studies. However, almost nothing is known about the dental tissue proportions of European Pleistocene hominins, apart from Neanderthal populations. This study aims to assess the three-dimensional dental tissue proportions of permanent canines belonging to the extensive sample of hominin teeth at Sierra de Atapuerca (Spain) through the use of microtomographic techniques. Our results show that early and middle Pleistocene populations from Atapuerca exhibit large coronal and root dentine dimensions, as well as a thinly enamelled pattern, which has been traditionally considered an autapomorphic Neanderthal trait. Therefore, these results might support an early enamel thickness decrease which is already observed 800kyr ago in Homo antecessor and maintained in later groups such as Sima de los Huesos and Neanderthal populations during the middle Pleistocene. Résumé Les patrons de proportions de l’émail et de la dentine ont éveillé un intérêt considérable dans les études phylogénétiques. Cependant, on ne connaît encore presque rien sur les proportions des tissus dentaires des hominidés du Pléistocène européen, à l’exception de celles des populations néandertaliennes. La présente étude vise à évaluer les proportions tridimensionnelles des tissus dentaires des canines permanentes issues d’un échantillon exceptionnel de dents d’homininés d’Atapuerca (Espagne) en utilisant des techniques microtomographiques. Nos résultats montrent que les populations du Pléistocène moyen et inférieur d’Atapuerca possèdent des proportions de dentine radiculaire et coronaire élevées, ainsi qu’un patron d’émail fin qui a été traditionnellement considéré comme un caractère autapomorphique des Néandertaliens. Par conséquent, ces résultats appuient l’apparition précoce d’une diminution de l’épaisseur de l’émail depuis au moins 800 ka chez Homo antecessor. Ce patron d’émail fin est également observé dans d’autres groupes plus tardifs, tels que Sima de los Huesos et les populations néandertaliennes pendant le Pléistocène moyen.

Modern humans sex estimation through dental tissue patterns of maxillary canines

Objectives Dental tissue proportions of human permanent canines is one of only a few sexually dimorphic features that is present in childhood and maintained in adults, offering the opportunity for this to be used in sex determination. This study assesses dental tissue volumes and surface areas of maxillary permanent canines in a sample of known sex to provide new data and to explore the potential of these variables as reliable sexual estimators. Materials and methods The teeth studied here derive from 56 individuals (27 females and 29 males) of known sex and age, and of different geographic origins. The teeth were scanned and three-dimensional (3D) measurements (volumes and surface areas) were obtained. In addition, a discriminant function analysis was applied. Results The results presented here concur with those previously published in relation to both size and dental tissue patterns. Male maxillary canines have a greater dentine component, whereas female enamel is thicker, leading to a difference in dental size in favor of males. Discriminant functions were calculated using these histological variables successfully identifying sex in between 87.5% and 93.75% of the known-sex hold-out sample, with 92.3% correctly assigned when all functions were applied together. Discussion The present study supports that methods for sex determination based on dental tissue measurements can achieve high allocation accuracies, being especially useful in the case of subadults or when no other appropriate method is available.

Tooth crown tissue proportions and enamel thickness in Early Pleistocene Homo antecessor molars (Atapuerca, Spain)

Tooth crown tissue proportions and enamel thickness distribution are considered reliable characters for inferring taxonomic identity, phylogenetic relationships, dietary and behavioural adaptations in fossil and extant hominids. While most Pleistocene hominins display variations from thick to hyper-thick enamel, Neanderthals exhibit relatively thinner. However, the chronological and geographical origin for the appearance of this typical Neanderthal condition is still unknown. The European late Early Pleistocene species Homo antecessor (Gran Dolina-TD6 site, Sierra de Atapuerca) represents an opportunity to investigate the appearance of the thin condition in the fossil record. In this study, we aim to test the hypothesis if H. antecessor molars approximates the Neanderthal condition for tissue proportions and enamel thickness. To do so, for the first time we characterised the molar inner structural organization in this Early Pleistocene hominin taxon (n = 17) and compared it to extinct and extant populations of the genus Homo from African, Asian and European origin (n = 355). The comparative sample includes maxillary and mandibular molars belonging to H. erectus, East and North African Homo, European Middle Pleistocene Homo, Neanderthals, and fossil and extant H. sapiens. We used high-resolution images to investigate the endostructural configuration of TD6 molars (tissue proportions, enamel thickness and distribution). TD6 permanent molars tend to exhibit on average thick absolute and relative enamel in 2D and 3D estimates, both in the complete crown and the lateral enamel. This condition is shared with the majority of extinct and extant hominin sample, except for Neanderthals and some isolated specimens. However, while the total crown percentage of dentine in TD6 globally resembles the low modern values, the lateral crown percentage of dentine tends to be much higher, closer to the Neanderthal signal. Similarly, the H. antecessor molar enamel distribution maps reveal a relative distribution pattern that is more similar to the Neanderthal condition (with the thickest enamel more spread at the periphery of the occlusal basin) rather than that of other fossil specimens and modern humans (with thicker cuspal enamel). Future studies on European Middle Pleistocene populations will provide more insights into the evolutionary trajectory of the typical Neanderthal dental structural organization.

Contribution of dental tissues to sex determination in modern human populations

Objectives Accurate sex estimation is an essential step for the reconstruction of the biological profile of human remains. Earlier studies have shown that elements of the human permanent dentition are sexually dimorphic. The aims of this study are to determine the degree of sexual dimorphism in the dental tissue volumes and surface areas of mandibular canines and to explore its potential for reliable sex determination. Method The teeth included in this study (n = 69) were selected from anthropological collections from Spain, South Africa and Sudan. In all cases, the sex of the individuals was known. The teeth were scanned and three-dimensional (3D) measurements (volumes and surfaces areas) were obtained. Finally, a dsicriminant function analysis was applied. Results Our results showed that sexual dimorphism in canine size is due to males having greater amounts of dentine, whereas enamel volume does not contribute significantly to overall tooth size dimorphism. Classification accuracy of the multivariable equations tested on slightly worn teeth ranged from 78 to 90.2% for the crossvalidation, and from 71.43 to 84.62% for the hold-out sample validation. When all functions were applied together, the sex was correctly assigned 92.30% of the time. Conclusions Our results suggest that the 3D variables from mandibular canine dental tissues are useful for sex determination as they present a high degree of dimorphism. The results obtained show the importance of 3D dental tissue measurements as a methodology in sex determination, which application should be considered as a supplemental method to others.

Metric and morphological comparison between the Arago (France) and Atapuerca-Sima de los Huesos (Spain) dental samples, and the origin of Neanderthals

The variability observed in the growing Middle Pleistocene hominin fossil record of Europe continues to trigger much debate on taxonomic issues and the biological processes that gave rise to Neanderthals. Here we present a metric and morphological comparative study of the dental samples recovered from the sites of Arago (southeast France) and Sima de los Huesos (SH) in the Sierra de Atapuerca (northern Spain). These sites are key to providing answers to these questions since they have yielded the largest hominin samples so far recovered for this time period. Despite the geographical proximity of the two sites and the contemporaneity of their hominin assemblages, we have observed remarkable metric and morphological differences between the teeth at Arago and SH. Whereas the SH teeth present an almost morphological identity with European Neanderthals, the Arago teeth exhibit a combination of plesiomorphic as well as some Neanderthal-derived features. In addition, the Arago crown dimensions are remarkably larger than those from SH, the differences being statistically significant for most variables. We hypothesize that during the Middle Pleistocene the European continent was settled at different points in time by hominin groups coming from Southwest Asia, probably from a common mother population evolving in this latter region. These first settlers can be identified by their more plesiomorphic morphology, whereas the most recent settlers are closer in appearance to Neanderthals. In addition, genetic processes such as isolation, genetic drift, directional adaptation or hybridization would have given rise to the puzzle we observe in the current fossil record.

A reassessment of the Montmaurin-La Niche mandible (Haute Garonne, France) in the context of European Pleistocene human evolution

We here present a comparative study of the Montmaurin-LN Middle Pleistocene mandible (Haute-Garonne, France). This mandible, of which its right and left molar series are preserved in situ, was found in La Niche cave (Montmaurin’s karst system) in 1949, and was first attributed to the ‘Mindel-Riss’ interglacial (= MIS 9 to 11) based on its geological context. Later studies based on geological and faunal evidence have attributed the Montmaurin-LN mandible to MIS 7. Following a detailed morphological and metric comparative study of the mandible in the 1970s, it was interpreted in the light of a still limited fossil record and the prevailing paradigm back then. Waiting for geochronological studies in the forthcoming years, here we review the main morphological and metrical features of this mandible and its molars, which have been reassessed in the framework of a remarkably enlarged Pleistocene fossil record since the mandible was first described, and our current, more in-depth understanding of human evolution in Europe. Using a selection of mandibular features with potential taxonomic signal we have found that the Montmaurin-LN mandible shares only a few derived traits with Neandertals. Our analyses reveal that this mandible is more closely related to the ancient specimens from the African and Eurasian Early and Middle Pleistocene, particularly due to the presence of primitive features of the Homo clade. In contrast, the external morphology of the molars is clearly similar to that of Neandertals. The results are assessed in the light of the present competing hypotheses used to explain the European hominin fossil record.

New methodology to reconstruct in 2-D the cuspal enamel of modern human lower molars

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.

Early Pleistocene hominin deciduous teeth from the Homo antecessor Gran Dolina-TD6 bearing level (Sierra de Atapuerca, Spain)

Objectives. During the last 13 years, the late Early Pleistocene Gran Dolina-TD6-2 level (Sierra de Atapuerca, northern Spain) has yielded an additional sample of 26 dental specimens attributed to Homo antecessor. In this report, we present a descriptive and comparative study of the six deciduous teeth. Methods. We provide external and internal morphological descriptions following classical terminology, as well as the mesiodistal and buccolingual measurements of the teeth. The internal morphology was described by means of micro-CT technique. Results. The TD6 deciduous teeth preserve primitive features regarding the Homo clade, such as the presence of styles in lower and upper canines and developed anterior and posterior foveae in the dm2. However, other features related to the complexity of the crown morphology (e.g., cingulum) are not present in this sample. Furthermore, the great reduction of the talonid of the dm1s is also noteworthy. Despite the limited comparative evidence, the presence of a remarkably well-developed tuberculum molare in the dm1 and dm1s from TD6 can be also considered a derived feature in the genus Homo. The TD6 hominins exhibit dental dimensions similar to those of other Pleistocene hominins. The dm1s are buccolingually elongated and the buccolingual diameter of ATD6-93 is the largest recorded so far in the Homo fossil record. Conclusions. This study expands the list of plesiomorphic features of H. antecessor, and provides some information on the evolutionary status of this species. However, the identification of some advanced traits evinces a step towards the derived morphology of European Pleistocene teeth. The study of the deciduous dentition confirms the mosaic pattern of H. antecessor morphology revealed in previous studies of this hominin sample.