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Abstract: The Pech-de-l''Azé I skull and mandible are included in the juvenile Neandertal remains from Europe. However, some preserved features in the cranial skeleton seem to distinguish the specimen from other Neandertal children. Unfortunately, the stratigraphic position and dating of this child has never been clear. Our recent work on unpublished archives show that the Pech-de-l''Azé I Neandertal child was discovered at the bottom of layer 6, attributed to the Mousterian of Acheulean tradition type B. These skull and mandible are the first diagnostic human remains (aside from an isolated tooth) attributed to the Mousterian of Acheulian tradition (MTA) type B. Consequently, we confirm that Neandertals were the makers of this Mousterian industry, which is characterized by unusual high frequencies of Upper Paleolithic type tools, elongated blanks and blades. We were able to date the context of the hominid remains by dating layer 6 and the layers above and beneath it using ESR, coupled ESR/230Th/234U (coupled ESR/U-series), and AMS 14C. Coupled ESR/U-series results on 16 mammalian teeth constrain the age of the uppermost layer 7 to 41–58ka, and layer 6 to 37–51ka. The wide spread in each age estimate results mainly from uncertainties in the gamma-dose rate. These ages are concordant with AMS 14C ages of two bones coming from the top of layer 6, which provide dates of about 41.7–43.6ka cal BP. A combination of stratigraphic arguments and dating results for layers 6 and 7 show that the Neandertal child cannot be older than 51ka or younger than 41ka. The lowermost layer 4 is shown to be older than 43ka by the principle of superposition and ESR dating in the immediately overlying layer 5. This study shows that the MTA type B had been manufactured by Neandertals before the arrival of anatomically modern humans in the local region. Additionally, by providing a firm chronological framework for the specific morphometric the features of Pech-de-l''Azé I Neandertal child, this study is a new step toward the understanding of temporal and spatial changes in the ontogenesis of Neandertals in south-western Europe during oxygen isotope stages 5–3. [Copyright &y& Elsevier]

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 δ 44/42 Ca 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., Competing Interests: Declaration of competing interest The authors declare no competing interests., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Competing Interests: Declaration of competing interest The authors declare no conflict interests.

The calcium isotopic composition (δ 44/42 Ca) of bone and tooth enamel can be used for dietary reconstructions of extant and extinct mammals. In natural conditions, the δ 44/42 Ca 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 δ 44/42 Ca values than carnivores. All the δ 44/42 Ca 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 δ 44/42 Ca 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 δ 44/42 Ca values. Mass balance calculations indicate that the low δ 44/42 Ca 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., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The understanding of Neanderthal societies, both with regard to their funerary behaviors and their subsistence activities, is hotly debated. Old excavations and a lack of taphonomic context are often factors that limit our ability to address these questions. To better appreciate the exact nature of what is potentially the oldest burial in Western Europe, Regourdou (Montignac-sur-Vézère, Dordogne), and to better understand the taphonomy of this site excavated more than 50 years ago, we report in this contribution a study of the most abundant animals throughout its stratigraphy: the European rabbit (Oryctolagus cuniculus). In addition to questions surrounding the potential bioturbation of the site's stratigraphy, analysis of the Regourdou rabbits could provide new information on Neandertal subsistence behavior. The mortality profile, skeletal-part representation, breakage patterns, surface modification, and comparison with modern reference collections supports the hypothesis that the Regourdou rabbit remains were primarily accumulated due to natural (attritional) mortality. Radiocarbon dates performed directly on the rabbit remains give ages ranging within the second half of Marine Isotope Stage 3, notably younger than the regional Mousterian period. We posit that rabbits dug their burrows within Regourdou's sedimentological filling, likely inhabiting the site after it was filled. The impact of rabbit activity now brings into question both the reliability of the archaeostratigraphy of the site and the paleoenvironmental reconstructions previously proposed for it, and suggests rabbits may have played a role in the distribution of the Neandertal skeletal remains., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

We present a bioanthropological study of dental remains recovered from El Salt Middle Palaeolithic site (Alcoy, Alicante, Spain). The dental remains were found in a sedimentary layer representing a calm depositional environment within a freshwater spring system. The corresponding archaeological context comprises a Middle Palaeolithic faunal and lithic assemblage that represents the last documented evidence of human occupation at the site, dating to between 47.2 ± 4.4 and 45.2 ± 3.4 ka (thousands of years ago). This evidence is overlain by an archaeologically sterile deposit dated to 44.7 ± 3.2 ka. Results show that the teeth belong to a single juvenile or young adult individual with morphological and metric features falling within the Neanderthal range of variability, although the considered traits are not taxonomically highly discriminant. The reported fossils are representative of the latest Middle Palaeolithic groups in the region and may be considered in the ongoing debate on the disappearance of Neanderthals and the end of the Middle Palaeolithic., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The globular braincase of modern humans is distinct from all fossil human species, including our closest extinct relatives, the Neandertals. Such adult shape differences must ultimately be rooted in different developmental patterns, but it is unclear at which point during ontogeny these group characteristics emerge. Here we compared internal shape changes of the braincase from birth to adulthood in Neandertals (N = 10), modern humans (N = 62), and chimpanzees (N = 62). Incomplete fossil specimens, including the two Neandertal newborns from Le Moustier 2 and Mezmaiskaya, were reconstructed using reference-based estimation methods. We used 3D geometric morphometrics to statistically compare shapes of virtual endocasts extracted from computed-tomographic scans. Throughout the analysis, we kept track of possible uncertainties due to the missing data values and small fossil sample sizes. We find that some aspects of endocranial development are shared by the three species. However, in the first year of life, modern humans depart from this presumably ancestral pattern of development. Newborn Neandertals and newborn modern humans have elongated braincases, and similar endocranial volumes. During a 'globularization-phase' modern human endocasts change to the globular shape that is characteristic for Homo sapiens. This phase of early development is unique to modern humans, and absent from chimpanzees and Neandertals. Our results support the notion that Neandertals and modern humans reach comparable adult brain sizes via different developmental pathways. The differences between these two human groups are most prominent directly after birth, a critical phase for cognitive development., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Spy cave (Jemeppe-sur-Sambre, Belgium) is reputed for the two adult Neandertal individuals discovered in situ in 1886. Recent reassessment of the Spy collections has allowed direct radiocarbon dating of these individuals. The sorting of all of the faunal collections has also led to the discovery of the remains of a Neandertal child, Spy VI. This individual is represented by two mandibular corpus fragments. The left fragment is the most complete and both sides preserve the mental foramen. Four deciduous teeth are associated with these mandibular remains: three incisors and one canine. The lower left canine (Spy 645a) conjoins with the corresponding alveolar socket in the left part of the mandible. Following extant standards, the developmental stage of the preserved teeth indicate an age at death of about one and a half years. In addition to performing a classical morphometric comparative study of the mandible and teeth, we have evaluated the dental tissue proportions using high-resolution microtomographic techniques. Our results show that Spy VI generally falls within the Neandertal range of variation. However, this specimen also exhibits particular traits, notably in the dental internal structural organization, which reveals that variation in the immature Neandertal variation is larger than what was variation currently represented by the available fossil record. These observations demonstrate the need for investigating the frequency and expression of immature Neandertal traits in fossil anterior teeth, as well as their temporal and geographic variation. Direct radiocarbon dating of the Spy VI specimen has been conducted in two different laboratories. The results of Spy VI confirm the age previously determined for the two adults, making the Spy Neandertal remains the youngest ever directly dated in northwest Europe., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

This comparative study of maxillae in Neandertals, Qafzeh, and extant children examines two specific traits: the premaxillary suture (sutura incisiva) and the interincisive sinuses, proposing a new hypothesis about some features of the Neandertal mid-face. Morphologic study of the premaxillary suture at its different borders (i.e. the nasal aspect of the frontal process, nasal and palatal aspects of the palatal process of the maxilla) indicates a persistence of the suture among very young Neandertal children in comparison to the condition in extant ones. This suggests a longer independence of some parts of the premaxilla in Neandertals. To further examine this possibility, CT scans of two Neandertal children were analyzed: Roc de Marsal, estimated to be about 3 years, and Engis 2, estimated to be about 5-6 years. The results are quite different between the fossils. In the older, the premaxillary suture is represented only by a deep groove. In the younger it extends deep to the surface of the nasal process reaching the Parinaud's canal. Synostosis of the premaxillary suture was found to occur later in Neandertal children than in modern ones. Moreover, we observed the existence of two interincisive sinuses in the fossil children, whereas this is rare in modern children (present on only 2% of our sample of 0-6 year-old infants, n = 247). Persistence of an open premaxillary suture represents the potential for an extended period of growth of the Neandertal mid-face. Although no trace of the premaxillary suture remains in adult Neandertals, Neandertals present many features classically considered as consequences of this persistence. The two interincisive sinuses could be a consequence of the labio-lingual diameter of the incisors. The results presented here can be further investigated by additional studies on the cranial sutural system and by precise morphologic observations and CT scans of the mid-face of a larger sample of fossil children.