Dental Paleobiology in a Juvenile Neanderthal (Combe-Grenal, Southwestern France).

Combe-Grenal site (Southwest France) was excavated by F. Bordes between 1953 and 1965. He found several human remains in Mousterian levels 60, 39, 35 and especially 25, corresponding to MIS 4 (~75-70/60 ky BP) and with Quina Mousterian lithics. One of the fossils found in level 25 is Combe-Grenal IV, consisting of a fragment of the left corpus of a juvenile mandible. This fragment displays initial juvenile periodontitis, and the two preserved teeth (LLP4 and LLM1) show moderate attrition and dental calculus. The SEM tartar analysis demonstrates the presence of cocci and filamentous types of bacteria, the former being more prevalent. This result is quite different from those obtained for the two adult Neanderthals Kebara 2 and Subalyuk 1, where more filamentous bacteria appear, especially in the Subalyuk 1 sample from Central Europe. These findings agree with the available biomedical data on periodontitis and tartar development in extant individuals, despite the different environmental conditions and diets documented by numerous archeological, taphonomical and geological data available on Neanderthals and present-day populations. New metagenomic analyses are extending this information, and despite the inherent difficulties, they will open important perspectives in studying this ancient human pathology.

On holes and strings: Earliest displays of human adornment in the Middle Palaeolithic.

Glycymeris shell beads found in Middle Palaeolithic sites are understood to be artifacts collected by modern humans for symbolic use. In Misliya Cave, Israel, dated to 240-160 ka BP, Glycymeris shells were found that were neither perforated nor manipulated; nevertheless, transportation to the cave is regarded as symbolic. In about 120 ka BP at Qafzeh Cave, Israel, modern humans collected naturally perforated Glycymeris shells also for symbolic use. Use-wear analyses backed by experiments demonstrate that the Qafzeh shells were suspended on string, thus suggesting that the collection of perforated shells was intentional. The older Misliya shells join a similar finding from South Africa, while the later-dated perforated shells from Qafzeh resemble other assemblages from North Africa and the Levant, also dated to about 120 ka BP. We conclude that between 160 ka BP and 120 ka BP there was a shift from collecting complete valves to perforated ones, which reflects both the desire and the technological ability to suspend shell beads on string to be displayed on the human body.

The Neanderthal teeth from Marillac (Charente, Southwestern France): Morphology, comparisons and paleobiology.

Few European sites have yielded human dental remains safely dated to the end of MIS 4/beginning of MIS 3. One of those sites is Marillac (Southwestern France), a collapsed karstic cave where archeological excavations (1967-1980) conducted by B. Vandermeersch unearthed numerous faunal and human remains, as well as a few Mousterian Quina tools. The Marillac sinkhole was occasionally used by humans to process the carcasses of different prey, but there is no evidence for a residential use of the site, nor have any hearths been found. Rare carnivore bones were also discovered, demonstrating that the sinkhole was seasonally used, not only by Neanderthals, but also by predators across several millennia. The lithostratigraphic units containing the human remains were dated to ∼60 kyr. The fossils consisted of numerous fragments of skulls and jaws, isolated teeth and several post-cranial bones, many of them with traces of perimortem manipulations. For those already published, their morphological characteristics and chronostratigraphic context allowed their attribution to Neanderthals. This paper analyzes sixteen unpublished human teeth (fourteen permanent and two deciduous) by investigating the external morphology and metrical variation with respect to other Neanderthal remains and a sample from modern populations. We also investigate their enamel thickness distribution in 2D and 3D, the enamel-dentine junction morphology (using geometric morphometrics) of one molar and two premolars, the roots and the possible expression of taurodontism, as well as pathologies and developmental defects. The anterior tooth use and paramasticatory activities are also discussed. Morphological and structural alterations were found on several teeth, and interpreted in light of human behavior (tooth-pick) and carnivores' actions (partial digestion). The data are interpreted in the context of the available information for the Eurasian Neanderthals.

From number sense to number symbols. An archaeological perspective.

How and when did hominins move from the numerical cognition that we share with the rest of the animal world to number symbols? Objects with sequential markings have been used to store and retrieve numerical information since the beginning of the European Upper Palaeolithic (42 ka). An increase in the number of markings and complexity of coding is observed towards the end of this period. The application of new analytical techniques to a 44-42 ka old notched baboon fibula from Border Cave, South Africa, shows that notches were added to this bone at different times, suggesting that devices to store numerical information were in use before the Upper Palaeolithic. Analysis of a set of incisions on a 72-60 ka old hyena femur from the Les Pradelles Mousterian site, France, indicates, by comparison with markings produced by modern subjects under similar constraints, that the incisions on the Les Pradelles bone may have been produced to record, in a single session, homologous units of numerical information. This finding supports the view that numerical notations were in use among archaic hominins. Based on these findings, a testable five-stage scenario is proposed to establish how prehistoric cultures have moved from number sense to the use of number symbols.This article is part of a discussion meeting issue 'The origins of numerical abilities'.

Earliest cranio-encephalic trauma from the Levantine Middle Palaeolithic: 3D reappraisal of the Qafzeh 11 skull, consequences of pediatric brain damage on individual life condition and social care.

The Qafzeh site (Lower Galilee, Israel) has yielded the largest Levantine hominin collection from Middle Palaeolithic layers which were dated to circa 90-100 kyrs BP or to marine isotope stage 5b-c. Within the hominin sample, Qafzeh 11, circa 12-13 yrs old at death, presents a skull lesion previously attributed to a healed trauma. Three dimensional imaging methods allowed us to better explore this lesion which appeared as being a frontal bone depressed fracture, associated with brain damage. Furthermore the endocranial volume, smaller than expected for dental age, supports the hypothesis of a growth delay due to traumatic brain injury. This trauma did not affect the typical human brain morphology pattern of the right frontal and left occipital petalia. It is highly probable that this young individual suffered from personality and neurological troubles directly related to focal cerebral damage. Interestingly this young individual benefited of a unique funerary practice among the south-western Asian burials dated to Middle Palaeolithic.

Neanderthal infant and adult infracranial remains from Marillac (Charente, France).

At the site of Marillac, near the Ligonne River in Marillac-le-Franc (Charente, France), a remarkable stratigraphic sequence has yielded a wealth of archaeological information, palaeoenvironmental data, as well as faunal and human remains. Marillac must have been a sinkhole used by Neanderthal groups as a hunting camp during MIS 4 (TL date 57,600 ± 4,600BP), where Quina Mousterian lithics and fragmented bones of reindeer predominate. This article describes three infracranial skeleton fragments. Two of them are from adults and consist of the incomplete shafts of a right radius (Marillac 24) and a left fibula (Marillac 26). The third fragment is the diaphysis of the right femur of an immature individual (Marillac 25), the size and shape of which resembles those from Teshik-Tash and could be assigned to a child of a similar age. The three fossils have been compared with the remains of other Neanderthals or anatomically Modern Humans (AMH). Furthermore, the comparison of the infantile femora, Marillac 25 and Teshik-Tash, with the remains of several European children from the early Middle Ages clearly demonstrates the robustness and rounded shape of both Neanderthal diaphyses. Evidence of peri-mortem manipulations have been identified on all three bones, with spiral fractures, percussion pits and, in the case of the radius and femur, unquestionable cutmarks made with flint implements, probably during defleshing. Traces of periostosis appear on the fibula fragment and on the immature femoral diaphysis, although their aetiology remains unknown.

Hyperostosis frontalis interna in a Neandertal from Marillac (Charente, France).

The site of Marillac (Charente, France) has yielded an important stratigraphic sequence containing numerous Neandertal remains (some of them with peri-mortem manipulations) from lithofacies 2 (Quina Mousterian). This level has been correlated with MIS 4 and is associated with a TL date of 57,600 ± 4600 years BP (before present). The study of one of the cranial fragments (Marillac 3) revealed a grade 2 or Type B Hyperostosis frontalis interna (HFI), remodelling and altering the internal table of the thick frontal bone. This pathology has been analysed macroscopically together with radiography and sections made using a microscanner and a scanner. The development of the HFI is compared with published evidence for Sangiran 3 (Homo erectus), two other Neandertals (Forbes' Quarry and Shanidar 5), and several archaeological samples. Forbes' Quarry seems to display more advanced HFI than either Shanidar 5 or Marillac 3. The three Neandertals may be considered mature individuals (≥40 years) and it seems likely that the aetiology of this pathology may be associated with hormonal alterations, as has been suggested for past and extant populations. While the prevalence of HFI in contemporary post-menopausal women is well documented, the identification of HFI amongst males from several archaeological samples (Neanderthals, Ancient Egypt, Syrian Bronze Age or the Anasazi), with different stages of development, confirm that the pathology affected both sexes in past populations. Additional data and research are still needed to elucidate the etiopathogenesis of this illness and to better understand the relationship between environmental factors and their possible influences/consequences for the development of metabolic disorders in prehistoric populations.

Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian.

The transition from the Middle Paleolithic (MP) to Upper Paleolithic (UP) is marked by the replacement of late Neandertals by modern humans in Europe between 50,000 and 40,000 y ago. Châtelperronian (CP) artifact assemblages found in central France and northern Spain date to this time period. So far, it is the only such assemblage type that has yielded Neandertal remains directly associated with UP style artifacts. CP assemblages also include body ornaments, otherwise virtually unknown in the Neandertal world. However, it has been argued that instead of the CP being manufactured by Neandertals, site formation processes and layer admixture resulted in the chance association of Neanderthal remains, CP assemblages, and body ornaments. Here, we report a series of accelerator mass spectrometry radiocarbon dates on ultrafiltered bone collagen extracted from 40 well-preserved bone fragments from the late Mousterian, CP, and Protoaurignacian layers at the Grotte du Renne site (at Arcy-sur-Cure, France). Our radiocarbon results are inconsistent with the admixture hypothesis. Further, we report a direct date on the Neandertal CP skeleton from Saint-Césaire (France). This date corroborates the assignment of CP assemblages to the latest Neandertals of western Europe. Importantly, our results establish that the production of body ornaments in the CP postdates the arrival of modern humans in neighboring regions of Europe. This new behavior could therefore have been the result of cultural diffusion from modern to Neandertal groups.

Shells and ochre in Middle Paleolithic Qafzeh Cave, Israel: indications for modern behavior.

Qafzeh Cave, the burial grounds of several anatomically modern humans, producers of Mousterian industry, yielded archaeological evidence reflecting their modern behavior. Dated to 92ka BP, the lower layers at the site contained a series of hearths, several human graves, flint artifacts, animal bones, a collection of sea shells, lumps of red ochre, and an incised cortical flake. The marine shells were recovered from layers earlier than most of the graves except for one burial. The shells were collected and brought from the Mediterranean Sea shore some 35km away, and are complete Glycymeris bivalves, naturally perforated. Several valves bear traces of having been strung, and a few had ochre stains on them.

Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: review and use of a multi-source mixing model.

The carbon and nitrogen isotopic abundances of the collagen extracted from the Saint-Césaire I Neanderthal have been used to infer the dietary behaviour of this specimen. A review of previously published Neanderthal collagen isotopic signatures with the addition of 3 new collagen isotopic signatures from specimens from Les Pradelles allows us to compare the dietary habits of 5 Neanderthal specimens from OIS 3 and one specimen from OIS 5c. This comparison points to a trophic position as top predator in an open environment, with little variation through time and space. In addition, a comparison of the Saint-Césaire I Neanderthal with contemporaneous hyaenas has been performed using a multi-source mixing model, modified from Phillips and Gregg (2003, Oecologia 127, 171). It appears that the isotopic differences between the Neanderthal specimen and hyaenas can be accounted for by much lower amounts of reindeer and much higher amounts of woolly rhinoceros and woolly mammoth in the dietary input of the Neanderthal specimen than in that of hyaenas, with relatively similar contributions of bovinae, large deer and horse for both predators, a conclusion consistent with the zooarchaeological data. The high proportion of very large herbivores, such as woolly rhinoceros and woolly mammoth, in Neanderthal's diet compare to that of the scavenging hyaenas suggests that Neanderthals could not acquire these prey through scavenging. They probably had to hunt for proboscideans and rhinoceros. Such a prey selection could result from a long lasting dietary tradition in Europe.

Evidence for interpersonal violence in the St. Cesaire Neanderthal.

The St. Césaire 1 Neanderthal skeleton of a young adult individual is unique in its association with Châtelperronian artifacts from a level dated to ca. 36,000 years ago. Computer-tomographic imaging and computer-assisted reconstruction of the skull revealed a healed fracture in the cranial vault. When paleopathological and forensic diagnostic standards are applied, the bony scar bears direct evidence for the impact of a sharp implement, which was presumably directed toward the individual during an act of interpersonal violence. These findings add to the evidence that Neanderthals used implements not only for hunting and food processing, but also in other behavioral contexts. It is hypothesized that the high intra-group damage potential inherent to weapons might have represented a major factor during the evolution of hominid social behavior.