Diverse bone-calcium isotope compositions in Neandertals suggest different dietary strategies.

Zooarcheological and geochemical evidence suggests Neanderthals were top predators, but their adherence to a strictly carnivorous diet has been questioned. Recent studies have demonstrated the potential of calcium-stable isotopes to evaluate trophic and ecological relationships. Here, we measure the δ44/42Ca values in bone samples from Mousterian contexts at Grotte du Bison (Marine Isotope Stage 3, Yonne, France) and Regourdou (Marine Isotope Stage 5, Dordogne, France) in two new Neanderthal individuals, associated fauna, and living local plants. We use a Bayesian mixing model to estimate the dietary composition of these Neanderthal individuals, plus a third one already analyzed. The results reveal three distinct diets: a diet including accidental or voluntary consumption of bone-based food, an intermediate diet, and a diet without consumption of bone-based food. This finding is the first demonstration of diverse subsistence strategies among Neanderthals and as such, reconciles archaeological and geochemical dietary evidence.

Sex estimation of the adult Neandertal Regourdou 1 (Montignac, France): Implications for sexing human fossil remains.

Sex is a biological trait fundamental to the study of hominin fossils. Among the many questions that can be addressed are those related to taxonomy, biological variability, sexual dimorphism, paleoobstetrics, funerary selection, and paleodemography. While new methodologies such as paleogenomics or paleoproteomics can be used to determine sex, they have not been systematically applied to Pleistocene human remains due to their destructive nature. Therefore, we estimated sex from the coxal bone of the newly discovered pelvic remains of the Regourdou 1 Neandertal (Southwest France, MIS 5) based on morphological and metric data employing two methods that have been recently revised and shown to be reliable in multiple studies. Both methods calculate posterior probabilities of the estimate. The right coxal bone of Regourdou 1 was partially reconstructed providing additional traits for sex estimation. These methods were cross validated on 14 sufficiently preserved coxal bones of specimens from the Neandertal lineage. Our results show that the Regourdou 1 individual, whose postcranial skeleton is not robust, is a male, and that previous sex attributions of comparative Neandertal specimens are largely in agreement with those obtained here. Our results encourage additional morphological research of fossil hominins in order to develop a set of methods that are applicable, reliable, and reproducible.

Isotopic calcium biogeochemistry of MIS 5 fossil vertebrate bones: application to the study of the dietary reconstruction of Regourdou 1 Neandertal fossil.

The calcium isotopic composition (δ44/42Ca) of bone and tooth enamel can be used for dietary reconstructions of extant and extinct mammals. In natural conditions, the δ44/42Ca value of bone and teeth varies according to dietary intake with a constant isotopic offset of about -0.6‰. Owing to the poor conservation of collagen, carbon (C), and nitrogen (N) isotopic compositions of the Regourdou Mousterian site (MIS 5, Dordogne, France) previously failed to provide any paleodietary information. Therefore, to reconstruct the trophic chain, we have measured calcium (Ca) isotopes from fossil bone samples of the fauna from the Regourdou site, as well as from three bone samples of the Regourdou 1 Neandertal specimen. The results show a taxon-dependent patterning of the Ca isotopic compositions: herbivores generally have higher δ44/42Ca values than carnivores. All the δ44/42Ca values of Regourdou 1 are low (<-1.6‰), placing this specimen amid carnivores. Using a bone-muscle Ca isotopic offset determined on extant animals, we further show that the δ44/42Ca value of the Regourdou 1 diet, and that of most carnivores, cannot be accounted for by the consumption of meat only, as plants and meat have indistinguishable δ44/42Ca values. Mass balance calculations indicate that the low δ44/42Ca values of the Neandertal's carnivorous diet are explained by the ingestion of bone marrow containing as little as 1% trabecular bone. Our results show that the Regourdou 1 Neanderthal consumed a mixture of various herbivorous prey, as well as trabecular bone, which probably occurred when marrow was ingested, by accident or intentionally.

A case of marked bilateral asymmetry in the sacral alae of the Neandertal specimen Regourdou 1 (Périgord, France).

OBJECTIVES: A marked asymmetry was previously reported in the sacral alae and S1-L5 facets orientation of the Neandertal individual Regourdou 1. Here, we provide a detailed description and quantification of the morphology and degree of asymmetry of this sacrum. MATERIAL AND METHODS: Regourdou 1 was compared to a modern human sample composed of 24 females and 17 males, and to other Neandertal individuals. Both traditional and geometric morphometric analyses were used in order to quantify the degree of sacral asymmetry of Regourdou 1. RESULTS: The asymmetry of both sacral alae and facets orientation substantially exceeds directional and absolute asymmetry of the healthy modern sample. Regourdou 1 shows a considerably shorter right ala, which is absolutely and relatively outside of the modern and Neandertal variations. CONCLUSION: Regourdou 1 shows marked sacral asymmetry that probably originated in early ontogenetic development. An asymmetric sacrum reflects asymmetric load dissipation and could relate to other morphological abnormalities observed in the skeleton, especially the mild scoliosis of the spine and the asymmetry of the femoral diaphyses. Further investigation is necessary to elucidate the relationship between those morphologies as well as a potential impact on the life of the individual.

Neandertal foot remains from Regourdou 1 (Montignac-sur-Vézère, Dordogne, France).

Regourdou is a well-known Middle Paleolithic site which has yielded the fossil remains of a minimum of two Neandertal individuals. The first individual (Regourdou 1) is represented by a partial skeleton while the second one is represented by a calcaneus. The foot remains of Regourdou 1 have been used in a number of comparative studies, but to date a full description and comparison of all the foot remains from the Regourdou 1 Neandertal, coming from the old excavations and from the recent reanalysis of the faunal remains, does not exist. Here, we describe and comparatively assess the Regourdou 1 tarsals, metatarsals and phalanges. They display traits observed in other Neandertal feet, which are different from some traits of the Sima de los Huesos (Atapuerca) hominins and of Middle Paleolithic, Upper Paleolithic and recent modern humans. These Neandertal features are: a rectangular talar trochlea with a large lateral malleolar facet, a broad talar head, a broad calcaneus with a projecting sustentaculum tali, a wide and wedged navicular with a projecting medial tubercle, large and wide bases of the lateral metatarsals, and mediolaterally expanded and robust phalanges that also show hallux valgus in a strongly built hallux.

The costal skeleton of the Regourdou 1 Neandertal.

The morphology and size of the Neandertal thorax is a subject of growing interest due to its link to general aspects of body size and shape, including physiological aspects related to bioenergetics and activity budgets. However, the number of well-preserved adult Neandertal costal remains is still low. The recent finding of new additional costal remains from the Regourdou 1 (R1) skeleton has rendered this skeleton as one of the most complete Neandertal costal skeletons with a minimum of 18 ribs represented, five of which are complete or virtually complete. Here we describe for the first time all the rib remains from R1 and compare them to a large modern Euroamerican male sample as well as to other published Neandertal individuals. The costal skeleton of this individual shows significant metric and morphological differences from our modern human male comparative sample. The perceived differences include: dorsoventrally large 1st and 2nd ribs, 3rd ribs with a very closed dorsal curvature and large maximum diameters at the posterior angle, a large tubercle-iliocostal line distance in the 4th rib, thick shafts at the dorsal end of its 6th ribs, thick mid-shafts of the 8th ribs, large articular tubercles at the 9th ribs, and thick shafts of the 11th and 12th ribs. Here we also describe a new mesosternal fragment: the left lateral half of sternebral segments 4 and 5. This portion reveals that the mesosternum of R1 had a sternal foramen in its inferiormost preserved sternal segment and supports previous estimation of the total length of this mesosternum. The new costal remains from R1 support the view that Neandertals, when compared with modern humans, show a significantly different thorax, consistent with differences found in other anatomical regions such as the vertebral column and pelvis.

The multiple maternal legacy of the Late Iron Age group of Urville-Nacqueville (France, Normandy) documents a long-standing genetic contact zone in northwestern France.

The compilation of archaeological and genetic data for ancient European human groups has provided persuasive evidence for a complex series of migrations, population replacements and admixture until the Bronze Age. If the Bronze-to-Iron Age transition has been well documented archaeologically, ancient DNA (aDNA) remains rare for the latter period and does not precisely reflect the genetic diversity of European Celtic groups. In order to document the evolution of European communities, we analysed 45 individuals from the Late Iron Age (La Tène) Urville-Nacqueville necropolis in northwestern France, a region recognized as a major cultural contact zone between groups from both sides of the Channel. The characterization of 37 HVS-I mitochondrial sequences and 40 haplogroups provided the largest maternal gene pool yet recovered for the European Iron Age. First, descriptive analyses allowed us to demonstrate the presence of substantial amounts of steppe-related mitochondrial ancestry in the community, which is consistent with the expansion of Bell Beaker groups bearing an important steppe legacy in northwestern Europe at approximately 2500 BC. Second, maternal genetic affinities highlighted with Bronze Age groups from Great Britain and the Iberian Peninsula regions tends to support the idea that the continuous cultural exchanges documented archaeologically across the Channel and along the Atlantic coast (during and after the Bronze Age period) were accompanied by significant gene flow. Lastly, our results suggest a maternal genetic continuity between Bronze Age and Iron Age groups that would argue in favour of a cultural transition linked to progressive local economic changes rather than to a massive influx of allochthone groups. The palaeogenetic data gathered for the Urville-Nacqueville group constitute an important step in the biological characterization of European Iron age groups. Clearly, more numerous and diachronic aDNA data are needed to fully understand the complex relationship between the cultural and biological evolution of groups from the period.

Analytical evidence of enamel hypomineralisation on permanent and primary molars amongst past populations.

Molar Incisor Hypomineralisation (MIH) and Hypomineralised Second Primary Molars (HSPM) involve prevalent qualitative structural developmental anomalies of tooth enamel affecting the first permanent molars (and often incisors) and the second primary molars, respectively. These demarcated hypomineralised lesions of enamel manifest as white-cream or yellow-brown opacities, with possible post-eruptive localised loss of enamel. Aetiological hypotheses have involved contemporary life factors (i.e. environmental pollutant exposure or early childhood medications) in contrast to factors not limited to a specific time period (i.e. hypoxia at birth or genetic predisposition). Evidence of MIH in ancient populations would reinforce aetiological factors present for many centuries. By means of microtomographic and X-ray fluorescence analyses the present study provides evidence that (i) two archaeological specimens: "S407" (Sains-en-Gohelle, France, 12th-16th centuries) and "B335" (Beauvais, France, 15th-18th centuries) were MIH-affected, and (ii) one individual "S323" was affected by HSPM and MIH (Sains-en-Gohelle, France, 7th-11th centuries).

Diagnostic guide enabling distinction between taphonomic stains and enamel hypomineralisation in an archaeological context.

OBJECTIVE: Molar Incisor Hypomineralisation (MIH) is a structural anomaly that affects the quality of tooth enamel and has important consequences for oral health. The developmentally hypomineralised enamel has normal thickness and can range in colour from white to yellow or brown with or without surface breakdown. The possibility of finding MIH in 'ancient populations' could downplay several current aetiological hypotheses (e.g., dioxin derivatives, bisphenols, antibiotics) without excluding the possible multifactorial aspect of the anomaly. In an archaeological context, chemical elements contained in the burial ground can stain teeth yellow or brown and therefore might create a taphonomic bias. The purpose of the present study is to test a proposed diagnostic guide enabling determination of the pathological or taphonomic cause of enamel discolouration and defects that resemble MIH present on 'ancient teeth'. DESIGN: Two sample groups including MIH discoloration (n=12 teeth) from living patients, taphonomic discoloration (n=9 teeth) and unknown discoloration (n=2 teeth) from medieval specimens were tested. Three non-destructive methods-Raman spectroscopy, X-ray micro-computed tomography and X-ray fluorescence were utilised. RESULTS: Hypomineralised enamel has decreased mineral density (p<0.0001) and increased phosphate/ß-carbonate ratio (p<0.01) compared to normal enamel whereas relative concentrations of manganese, copper, iron and lead are similar. In taphonomic discoloration, relative concentrations of these elements are significantly different (p<0.05) to normal enamel whereas mineral density and Raman spectra profile are comparable. CONCLUSIONS: Enamel hypomineralisation can be distinguished from taphonomic staining in archaeological teeth.

Mineral density of hypomineralised and sound enamel.

Molar Incisor Hypomineralisation (MIH) is a structural anomaly that affects the quality of tooth enamel and has important consequences for oral health. The developmentally hypomineralised enamel has normal thickness and can range in colour from white to yellow or brown. The purpose of the present study is to compare the mineral density of hypomineralised and normal enamel. The sample included eight MIH teeth from seven patients. MIH teeth were scanned using high resolution microtomography. Non-parametric statistical tests (Wilcoxon test for paired samples) were carried out. Hypomineralised enamel has decreased mineral density (mean 19%; p < 0.0001) compared to normal enamel. This weak enamel has implications in clinical management of MIH lesions.

When the waves of European Neolithization met: first paleogenetic evidence from early farmers in the southern Paris Basin.

An intense debate concerning the nature and mode of Neolithic transition in Europe has long received much attention. Recent publications of paleogenetic analyses focusing on ancient European farmers from Central Europe or the Iberian Peninsula have greatly contributed to this debate, providing arguments in favor of major migrations accompanying European Neolithization and highlighting noticeable genetic differentiation between farmers associated with two archaeologically defined migration routes: the Danube valley and the Mediterranean Sea. The aim of the present study was to fill a gap with the first paleogenetic data of Neolithic settlers from a region (France) where the two great currents came into both direct and indirect contact with each other. To this end, we analyzed the Gurgy 'Les Noisats' group, an Early/Middle Neolithic necropolis in the southern part of the Paris Basin. Interestingly, the archaeological record from this region highlighted a clear cultural influence from the Danubian cultural sphere but also notes exchanges with the Mediterranean cultural area. To unravel the processes implied in these cultural exchanges, we analyzed 102 individuals and obtained the largest Neolithic mitochondrial gene pool so far (39 HVS-I mitochondrial sequences and haplogroups for 55 individuals) from a single archaeological site from the Early/Middle Neolithic period. Pairwise FST values, haplogroup frequencies and shared informative haplotypes were calculated and compared with ancient and modern European and Near Eastern populations. These descriptive analyses provided patterns resulting from different evolutionary scenarios; however, the archaeological data available for the region suggest that the Gurgy group was formed through equivalent genetic contributions of farmer descendants from the Danubian and Mediterranean Neolithization waves. However, these results, that would constitute the most ancient genetic evidence of admixture between farmers from both Central and Mediterranean migration routes in the European Neolithization debate, are subject to confirmation through appropriate model-based approaches.

The vertebral column of the Regourdou 1 Neandertal.

The Regourdou 1 partial skeleton was found in 1957 in level IV of the eponymous site located in Montignac-sur-Vézère (Dordogne, France) and until now it has been only partially published. The ongoing revision of the faunal remains from the site has yielded additional fossils that pertain to this skeleton. Here we study the vertebral column of this individual, providing for the first time detailed descriptions for all of the fossils and reassessing the anatomical position of all of the fragments. The vertebral column of Regourdou 1 is one of the most complete in the Neandertal fossil record with at least 20 pre-sacral vertebrae (seven cervicals, nine thoracic and four lumbars), a partial sacrum and a fragmentary first coccygeal vertebra. When compared with modern humans, the vertebrae of Regourdou 1 display significant metric differences, and fit well within the range of Neandertal variability. A preliminary analysis of the most complete thoracic vertebrae of this individual indicates that Neandertals displayed significant differences from modern humans in the thoracic spine, which adds to the differences already observed in the cervical and lumbar regions. Finally, we have also observed mild signs of osteoarthrosis, albeit to a lower degree of that present in other Neandertals such as La Chapelle-aux-Saints, La Ferrassie 1 or Shanidar 3. This is consistent with the younger adult age for Regourdou 1.

The mesosternum of the Regourdou 1 Neandertal revisited.

Fossil hominin mesosterna, while scarce, can provide useful morphological data in addition to rib remains regarding aspects of thoracic size and shape. These data, in turn, can address hypotheses related to respiratory dynamics, climatic adaptation, and ecogeographical patterning. In this study, we re-evaluate the anatomical representation of the mesosternum of the Regourdou 1 Neandertal individual that alters key aspects of the original description of the fossil remains. We compare this specimen together with the mesosterna of the Kebara 2 Neandertal male individual and the Tabun C1 Neandertal female individual to a large extant modern sample. Our study shows that the current evidence available for Neandertals indicates longer mesosterna, reflecting larger thorax sizes among Neandertals, in comparison with extant humans. Additionally, while this study weakens previous suggestions of ecogeographically mediated differences in the size and shape of upper thorax between Neandertals from the Mediterranean Levant and those deriving from Western Europe, we cannot unambiguously disprove the notion of such clinal differences.