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The use of 'teeth as tools' (non-masticatory or cultural-related dental wear) has largely been employed as a proxy for studying of past human behavior, mainly in permanent dentition from adult individuals. Here we present the analysis of the non-masticatory dental wear modifications on the deciduous dentition assigned to eight Neanderthal and anatomically modern human subadult individuals from Mousterian to Magdalenian technocultural contexts in the Cantabrian region (Northern Spain). Although preliminary, we tentatively suggest that these eight subadults present activity-related dental wear, including cultural striations, chipped enamel, toothpick grooves, and subvertical grooves. We also found evidence of habitual dental hygienic practices in the form of toothpicking on a deciduous premolar. Orientation of the cultural striations indicates similar handedness development as in modern children. Taken together, these dental wear patterns support the participation of young individuals in group activities, making them potential contributors to group welfare. This study potentially adds new evidence to the importance of the use of the mouth in paramasticatory activities or as a third hand throughout the Pleistocene, which can be confirmed with a more specific reference sample., Competing Interests: Declaration of competing interest The authors have no competing interests to declare., (Copyright © 2021 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Entheses have rarely been systematically studied in the field of human evolution. However, the investigation of their morphological variability (e.g., robusticity) could provide new insight into their evolutionary significance in the European Neanderthal populations. The aim of this work is to study the entheses and joint features of the lower limbs of El Sidrón Neanderthals (Spain; 49 ka), using standardized scoring methods developed on modern samples. Paleobiology, growth, and development of both juveniles and adults from El Sidrón are studied and compared with those of Krapina Neanderthals (Croatia, 130 ka) and extant humans. The morphological patterns of the gluteus maximus and vastus intermedius entheses in El Sidrón, Krapina, and modern humans differ from one another. Both Neanderthal groups show a definite enthesis design for the gluteus maximus, with little intrapopulation variability with respect to modern humans, who are characterized by a wider range of morphological variability. The gluteus maximus enthesis in the El Sidrón sample shows the osseous features of fibrous entheses, as in modern humans, whereas the Krapina sample shows the aspects of fibrocartilaginous ones. The morphology and anatomical pattern of this enthesis has already been established during growth in all three human groups. One of two and three of five adult femurs from El Sidrón and from Krapina, respectively, show the imprint of the vastus intermedius, which is absent among juveniles from those Neanderthal samples and in modern samples. The scant intrapopulation and the high interpopulation variability in the two Neanderthal samples is likely due to a long-term history of small, isolated populations with high levels of inbreeding, who also lived in different ecological conditions. The comparison of different anatomical entheseal patterns (fibrous vs. fibrocartilaginous) in the Neanderthals and modern humans provides additional elements in the discussion of their functional and genetic origin., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

The comment by DeSilva challenges our suggestion that brain growth of the El Sidrón J1 Neandertal was still incomplete at 7.7 years of age. Evidence suggests that endocranial volume is likely to represent less than 90% adult size at El Sidrón as well as Neandertal male plus Krapina samples, in line with further evidence from endocranial surface histology and dural sinus groove size., (Copyright © 2018, American Association for the Advancement of Science.)

Twenty-nine carpal bones of Homo neanderthalensis have been recovered from the site of El Sidrón (Asturias, Spain) during excavations between 1994 and 2009, alongside ∼2500 other Neandertal skeletal elements dated to ∼49,000 years ago. All bones of the wrist are represented, including adult scaphoids (n = 6), lunates (n = 2), triquetra (n = 4), pisiforms (n = 2), trapezia (n = 2), trapezoids (n = 5), capitates (n = 5), and hamates (n = 2), as well as one fragmentary and possibly juvenile scaphoid. Several of these carpals appear to belong to the complete right wrist of a single individual. Here we provide qualitative and quantitative morphological descriptions of these carpals, within a comparative context of other European and Near Eastern Neandertals, early and recent Homo sapiens, and other fossil hominins, including Homo antecessor, Homo naledi, and australopiths. Overall, the El Sidrón carpals show characteristics that typically distinguish Neandertals from H. sapiens, such as a relatively flat first metacarpal facet on the trapezium and a more laterally oriented second metacarpal facet on the capitate. However, there are some distinctive features of the El Sidrón carpals compared with most other Neandertals. For example, the tubercle of the trapezium is small with limited projection, while the scaphoid tubercle and hamate hamulus are among the largest seen in other Neandertals. Furthermore, three of the six adult scaphoids show a distinctive os-centrale portion, while another is a bipartite scaphoid with a truncated tubercle. The high frequency of rare carpal morphologies supports other evidence of a close genetic relationship among the Neandertals found at El Sidrón., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Fossil hominids often processed material held between their upper and lower teeth. Pulling with one hand and cutting with the other, they occasionally left impact cut marks on the lip (labial) surface of their incisors and canines. From these actions, it possible to determine the dominant hand used. The frequency of these oblique striations in an array of fossil hominins documents the typically modern pattern of 9 right- to 1 left-hander. This ratio among living Homo sapiens differs from that among chimpanzees and bonobos and more distant primate relatives. Together, all studies of living people affirm that dominant right-handedness is a uniquely modern human trait. The same pattern extends deep into our past. Thus far, the majority of inferred right-handed fossils come from Europe, but a single maxilla from a Homo habilis, OH-65, shows a predominance of right oblique scratches, thus extending right-handedness into the early Pleistocene of Africa. Other studies show right-handedness in more recent African, Chinese, and Levantine fossils, but the sample compiled for non-European fossil specimens remains small. Fossil specimens from Sima del los Huesos and a variety of European Neandertal sites are predominately right-handed. We argue the 9:1 handedness ratio in Neandertals and the earlier inhabitants of Europe constitutes evidence for a modern pattern of handedness well before the appearance of modern Homo sapiens., (© 2017 Wiley Periodicals, Inc.)

Objectives: The El Sidrón tali sample is assessed in an evolutionary framework. We aim to explore the relationship between Neandertal talus morphology and body size/shape. We test the hypothesis 1: talar Neandertal traits are influenced by body size, and the hypothesis 2: shape variables independent of body size correspond to inherited primitive features., Materials and Methods: We quantify 35 landmarks through 3D geometric morphometrics techniques to describe H. neanderthalensis-H. sapiens shape variation, by Mean Shape Comparisons, Principal Component, Phenetic Clusters, Minimum spanning tree analyses and partial least square and regression of talus shape on body variables. Shape variation correlated to body size is compared to Neandertals-Modern Humans (MH) evolutionary shape variation. The Neandertal sample is compared to early hominins., Results: Neandertal talus presents trochlear hypertrophy, a larger equality of trochlear rims, a shorter neck, a more expanded head, curvature and an anterior location of the medial malleolar facet, an expanded and projected lateral malleolar facet and laterally expanded posterior calcaneal facet compared to MH., Discussion: The Neandertal talocrural joint morphology is influenced by body size. The other Neandertal talus traits do not co-vary with it or not follow the same co-variation pattern as MH. Besides, the trochlear hypertrophy, the trochlear rims equality and the short neck could be inherited primitive features; the medial malleolar facet morphology could be an inherited primitive feature or a secondarily primitive trait; and the calcaneal posterior facet would be an autapomorphic feature of the Neandertal lineage., (© 2017 Wiley Periodicals, Inc.)

The study of the Neanderthal thorax has attracted the attention of the scientific community for more than a century. It is agreed that Neanderthals have a more capacious thorax than modern humans, but whether this was caused by a medio-lateral or an antero-posterior expansion of the thorax is still debated, and is key to understanding breathing biomechanics and body shape in Neanderthals. The fragile nature of ribs, the metameric structure of the thorax and difficulties in quantifying thorax morphology all contribute to uncertainty regarding precise aspects of Neanderthal thoracic shape. The El Sidrón site has yielded costal remains from the upper to the lower thorax, as well as several proximal rib ends (frequently missing in the Neanderthal record), which help to shed light on Neanderthal thorax shape. We compared the El Sidrón costal elements with ribs from recent modern humans as well as with fossil modern humans and other Neanderthals through traditional morphometric methods and 3D geometric morphometrics, combined with missing data estimation and virtual reconstruction (at the 1st, 5th and 11th costal levels). Our results show that Neanderthals have larger rib heads and articular tubercles than their modern human counterparts. Neanderthal 1st ribs are smaller than in modern humans, whereas 5th and 11th ribs are considerably larger. When we articulated mean ribs (size and shape) with their corresponding vertebral elements, we observed that compared to modern humans the Neanderthal thorax is medio-laterally expanded at every level, especially at T5 and T11. Therefore, in the light of evidence from the El Sidrón costal remains, we hypothesize that the volumetric expansion of the Neanderthal thorax proposed by previous authors would mainly be produced by a medio-lateral expansion of the thorax., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Ontogenetic studies help us understand the processes of evolutionary change. Previous studies on Neandertals have focused mainly on dental development and inferred an accelerated pace of general growth. We report on a juvenile partial skeleton (El Sidrón J1) preserving cranio-dental and postcranial remains. We used dental histology to estimate the age at death to be 7.7 years. Maturation of most elements fell within the expected range of modern humans at this age. The exceptions were the atlas and mid-thoracic vertebrae, which remained at the 5- to 6-year stage of development. Furthermore, endocranial features suggest that brain growth was not yet completed. The vertebral maturation pattern and extended brain growth most likely reflect Neandertal physiology and ontogenetic energy constraints rather than any fundamental difference in the overall pace of growth in this extinct human., (Copyright © 2017 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Well preserved thoracic vertebrae of Neandertals are rare. However, such fossils are important as their three-dimensional (3D) spatial configuration can contribute to the understanding of the size and shape of the thoracic spine and the entire thorax. This is because the vertebral body and transverse processes provide the articulation and attachment sites for the ribs. Dorsal orientation of the transverse processes relative to the vertebral body also rotates the attached ribs in a way that could affect thorax width. Previous research indicates possible evidence for greater dorsal orientation of the transverse processes and small vertebral body heights in Neandertals, but their 3D vertebral structure has not yet been addressed. Here we present 15 new vertebral remains from the El Sidrón Neandertals (Asturias, Northern Spain) and used 3D geometric morphometrics to address the above issues by comparing two particularly well preserved El Sidrón remains (SD-1619, SD-1641) with thoracic vertebrae from other Neandertals and a sample of anatomically modern humans. Centroid sizes of El Sidrón vertebrae are within the human range. Neandertals have larger T1 and probably also T2. The El Sidrón vertebrae are similar in 3D shape to those of other Neandertals, which differ from Homo sapiens particularly in central-lower regions (T6-T10) of the thoracic spine. Differences include more dorsally and cranially oriented transverse processes, less caudally oriented spinous processes, and vertebral bodies that are anteroposteriorly and craniocaudally short. The results fit with current reconstructions of Neandertal thorax morphology., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Objectives: During the microscopic examination of the Neandertal dentitions from El Sidrón (Spain) and Hortus (France), we found unusual fine parallel microstriations on the mesial and distal sides of all tooth types, near the cervix. As its appearance was similar to toothpick grooves described in other Homo species, it could correspond to early stages on its formation. To test this hypothesis we developed an experimental replication of a groove using grass stalks., Materials and Methods: Comparisons between 204 isolated Neandertal teeth and the two experimental dental specimens corroborate that the marks correspond to initial stages of toothpick groove formation, and we propose a five-grade recording scale that summarized the groove formation process., Results: Using this new recording procedure, we found that Hortus individuals have higher incidence of this trait (eight individuals out of nine) than the El Sidrón individuals (nine out of 11). Toothpick grooves from El Sidrón show the earliest stages of development, whereas the grooves found on Hortus Neandertals were well-developed. Toothpick grooves were also found in 21 incisors and canines., Conclusions: These differences could be due to the more advanced occlusal dental wear in Hortus individuals, maybe age-related and with a more meat-based diet maybe favoring the inclusion of food debris and thus probing as the cleaning methodology. Our results allow the identification and characterization of incipient toothpick grooves on the human fossil record and contribute to increase our knowledge on Neandertals behavioral and oral care habits., (© 2016 Wiley Periodicals, Inc.)

Here, we present the analysis of occlusal molar microwear textures of eight individuals from the El Sidrón Neandertal group (Spain). The aims of the study were: 1) to document potential age-, sex-, and maternal lineage-related differences in diet within a Neandertal familial group, and 2) to place the diet of El Sidrón individuals in the context of those of other Neandertal groups. This study also offers an interpretation of the diet of the El Sidrón Neandertals by comparing their microwear signatures to those of recent hunter-gatherer populations with diverse but known diets. The intra-group examination of the microwear signatures are consistent with the females of the El Sidrón group having had more abrasive diets or having used their teeth in more para-masticatory activities than did the males. Aside from the potential sex-related differences in diet, no additional intra-group dietary separation, such as by age group or maternal lineage, was observed. In comparison to other Neandertals, El Sidrón individuals, as a group, have microwear signatures most similar to those of other Neandertals from wooded habitats and different from those that lived in more open habitats. This result is expected based on the available paleoenvironmental reconstructions from El Sidrón Cave. The diet of the El Sidrón Neandertals, just like their Neandertal counterparts from similar wooded habitats, is interpreted as having been mixed, consisting of both meat and vegetable foods., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

We undertook a three-dimensional geometric morphometric (3DGM) analysis on 12 new Neandertal clavicle specimens from the El Sidrón site (Spain), dated to 49,000 years ago. The 3DGM methods were applied in a comparative framework in order to improve our understanding of trait polarity in features related to Homo pectoral girdle evolution, using other Neandertals, Homo sapiens, Pan, ATD6-50 (Homo antecessor), and KNM-WT 15000 (Homo ergaster/erectus) in the reference collection. Twenty-nine homologous landmarks were measured for each clavicle. Variation and morphological similarities were assessed through principal component analysis, conducted separately for the complete clavicle and the diaphysis. On average, Neandertal clavicles had significantly larger muscular entheses, double dorsal curvature, clavicle torsion, and cranial orientation of the acromial end than non-Neandertal clavicles; the El Sidrón clavicles fit this pattern. Variation within the samples was large, with extensive overlap between Homo species; only chimpanzee specimens clearly differed from the other specimens in morphometric terms. Taken together, our morphometric analyses are consistent with the following phylogenetic sequence. The primitive condition of the clavicle is manifest in the cranial orientation of both the acromial and sternal ends. The derived condition expressed in the H. sapiens + Neandertal clade is defined by caudal rotation of both the sternal and acromial ends, but with variation in the number of acromia remaining in a certain cranial orientation. Finally, the autapomorphic Neandertal condition is defined by secondarily acquired primitive cranial re-orientation of the acromial end, which varies from individual to individual. These results suggest that the pace of phylogenetic change in the pectoral girdle does not seem to follow that of other postcranial skeletal features., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

We present here the first cases in Neandertals of congenital clefts of the arch of the atlas. Two atlases from El Sidrón, northern Spain, present respectively a defect of the posterior (frequency in extant modern human populations ranging from 0.73% to 3.84%), and anterior (frequency in extant modern human populations ranging from 0.087% to 0.1%) arch, a condition in most cases not associated with any clinical manifestation. The fact that two out of three observable atlases present a low frequency congenital condition, together with previously reported evidence of retained deciduous mandibular canine in two out of ten dentitions from El Sidrón, supports the previous observation based on genetic evidence that these Neandertals constituted a group with close genetic relations. Some have proposed for humans and other species that the presence of skeletal congenital conditions, although without clinical significance, could be used as a signal of endogamy or inbreeding. In the present case this interpretation would fit the general scenario of high incidence of rare conditions among Pleistocene humans and the specific scenariothat emerges from Neandertal paleogenetics, which points to long-term small and decreasing population size with reduced and isolated groups. Adverse environmental factors affecting early pregnancies would constitute an alternative, non-exclusive, explanation for a high incidence of congenital conditions. Further support or rejection of these interpretations will come from new genetic and skeletal evidence from Neandertal remains.

A new collection of 49,000 year old Neandertal fossil humeri from the El Sidrón cave site (Asturias, Spain) is presented. A total of 49 humeral remains were recovered, representing 10 left and 8 right humeri from adults, adolescents, and a juvenile (not included in the analyses). 3D geometric morphometric (GM) methods as well as classic anthropological variables were employed to conduct a broad comparative analysis by means of mean centroid size and shape comparisons, principal components analysis, and cluster studies. Due to the fragmentary nature of the fossils, comparisons were organized in independent analyses according to different humeral portions: distal epiphysis, diaphysis, proximal epiphysis, and the complete humerus. From a multivariate viewpoint, 3D-GM analyses revealed major differences among taxonomic groups, supporting the value of the humerus in systematic classification. Notably, the Australopithecus anamensis (KP-271) and Homo ergaster Nariokotome (KNM-WT 15000) distal humerus consistently clusters close to those of modern humans, which may imply a primitive condition for Homo sapiens morphology. Australopithecus specimens show a high degree of dispersion in the morphospace. The El Sidrón sample perfectly fits into the classic Neandertal pattern, previously described as having a relatively wide olecranon fossa, as well as thin lateral and medial distodorsal pillars. These characteristics were also typical of the Sima de los Huesos (Atapuerca) sample, African mid-Pleistocene Bodo specimen, and Lower Pleistocene TD6-Atapuerca remains and may be considered as a derived state. Finally, we hypothesize that most of the features thought to be different between Neandertals and modern humans might be associated with structural differences in the pectoral girdle and shoulder joint., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Reconstructing the morphology of the Neanderthal rib cage not only provides information about the general evolution of human body shape but also aids understanding of functional anatomy and energetics. Despite this paleobiological importance there is still debate about the nature and extent of variations in the size and shape of the Neandertal thorax. The El Sidrón Neandertals can be used to contribute to this debate, providing new costal remains ranging from fully preserved and undistorted ribs to highly fragmented elements. Six first ribs are particularly well preserved and offer the opportunity to analyze thorax morphology in Neandertals. The aims of this paper are to present this new material, to compare the ontogenetic trajectories of the first ribs between Neandertals and modern humans, and, using geometric morphometrics, to test the hypothesis of morphological integration between the first rib and overall thorax morphology. The first ribs of the El Sidrón adult Neandertals are smaller in centroid size and tend to be less curved when compared with those of modern humans, but are similar to Kebara 2. Our results further show that the straightening of the first ribs is significantly correlated with a straightening of the ribs of the upper thorax (R = 0.66; p < 0.0001) in modern humans, suggesting modularity in the upper and lower thorax units as reported in other hominins. It also supports the hypothesis that the upper thorax of Neandertals differs in shape from modern humans with more anteriorly projecting upper ribs during inspiration. These differences could have biomechanical consequences and account for stronger muscle attachments in Neandertals. Different upper thorax shape would also imply a different spatial arrangement of the shoulder girdle and articulation with the humerus (torsion) and its connection to the upper thorax. Future research should address these inferences in the context of Neandertal overall body morphology., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The analysis of activity-related dental wear patterns in prehistoric anatomically modern humans and modern hunter-gatherers has shown sex differences attributable to a gendered division of labor. Neandertals are known to have extensive anterior dental wear related to the use of their front teeth as a tool. In this study we analyze the i) cultural striations (scratches on the labial surface of the anterior teeth with a cut-mark morphology), and ii) dental chipping (ante-mortem microfracture involving enamel or both enamel and dentine) in 19 Neandertal individuals from the l'Hortus (France), Spy (Belgium), and El Sidrón (Spain) sites, and compare the characteristics of those traits with the age and sex estimation for the individuals and among samples. The study reveals that all individuals have cultural striations, but those detected on the adult females are longer than the striations found in adult males. Regarding the distribution of dental chipping, the prevalence of this trait is higher in the maxillary dentition of males whereas females have the majority of dental chipping on their mandibular teeth. The differences detected on the overall activity-related dental wear pattern denote a difference or a division of labor by age and sex in Neandertals while using the mouth as a third hand, i.e., in activities other than the provisioning of food, and provide new evidence for the lifestyle of this Pleistocene fossil human species., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The developed cognitive capabilities for Homo sapiens seems to be the result of a specialized and lateralized brain, and as a result of this, humans display the highest degree of manual specialization or handedness among the primates. Studies regarding its emergence and distribution within the genus Homo show that handedness is present very early. The mode in which it was articulated and spread across the different species during the course of human evolution could provide information about our own cognitive capacities. Here we report the manual laterality attributed to eleven 49,000 old Neandertal individuals from El Sidrón cave (Spain), through the study of instrumental or cultural striations on the anterior dentition. Our results show a predominant pattern addressed to right-handers. These results fit within the modern human handedness distribution pattern and provide indirect evidence for behavior and brain lateralization on Neandertals. They support the early establishment of handedness in our genus. Moreover, the individual identified as Juvenile 1 (6-8 years old at death), displays the same striation pattern as the adult Neandertals from the sample, and thereby the ontogenic development of manual laterality in that Neandertal population seems to be similar to that of living modern humans.

Neanderthals disappeared sometime between 30,000 and 24,000 years ago. Until recently, Neanderthals were understood to have been predominantly meat-eaters; however, a growing body of evidence suggests their diet also included plants. We present the results of a study, in which sequential thermal desorption-gas chromatography-mass spectrometry (TD-GC-MS) and pyrolysis-gas chromatography-mass spectrometry (Py-GC-MS) were combined with morphological analysis of plant microfossils, to identify material entrapped in dental calculus from five Neanderthal individuals from the north Spanish site of El Sidrón. Our results provide the first molecular evidence for inhalation of wood-fire smoke and bitumen or oil shale and ingestion of a range of cooked plant foods. We also offer the first evidence for the use of medicinal plants by a Neanderthal individual. The varied use of plants that we have identified suggests that the Neanderthal occupants of El Sidrón had a sophisticated knowledge of their natural surroundings which included the ability to select and use certain plants.

Skull morphology results from the bone remodelling mechanism that underlies the specific bone growth dynamics. Histological study of the bone surface from Neanderthal mandible specimens of El Sidrón (Spain) provides information about the distribution of the remodelling fields (bone remodelling patterns or BRP) indicative of the bone growth directions. In comparison with other primate species, BRP shows that Neanderthal mandibles from the El Sidrón (Spain) sample present a specific BRP. The interpretation of this map allows inferences concerning the growth directions that explain specific morphological traits of the Neanderthal mandible, such as its quadrangular shape and the posterior location of the mental foramen., (This journal is © 2011 The Royal Society)

Neanderthals disappeared sometime between 30,000 and 24,000 years ago. Until recently, Neanderthals were understood to have been predominantly meat-eaters; however, a growing body of evidence suggests their diet also included plants. We present the results of a study, in which sequential thermal desorption-gas chromatography-mass spectrometry (TD-GC-MS) and pyrolysis-gas chromatography-mass spectrometry (Py-GC-MS) were combined with morphological analysis of plant microfossils, to identify material entrapped in dental calculus from five Neanderthal individuals from the north Spanish site of El Sidrón. Our results provide the first molecular evidence for inhalation of wood-fire smoke and bitumen or oil shale and ingestion of a range of cooked plant foods. We also offer the first evidence for the use of medicinal plants by a Neanderthal individual. The varied use of plants that we have identified suggests that the Neanderthal occupants of El Sidrón had a sophisticated knowledge of their natural surroundings which included the ability to select and use certain plants. [ABSTRACT FROM AUTHOR]

Abstract: This paper synthesizes and updates the information coming from the El Sidrón (Asturias, Northern Spain) neandertal site. Since 2000, a new sample of Homo neanderthalensis dated to at least 49,000years old is being systematically recovered at the El Sidrón cave site. The bone assemblage is located in a secondary position, and certainly derives from a close location. The sample is almost exclusively composed of human remains. There is a moderate number of Middle Paleolithic stone tools (n ≈415) and very few macro-faunal remains. All skeletal parts are preserved, including some rare bones such as the hyoid bone. Teeth are abundant (n =213), cranial and postcranial remains are also well represented, but fragmentary, with a special presence of foot and hand bones. A minimum number of thirteen individuals has been identified, comprising different developmental stages from infancy to adulthood: one infant, two juveniles, three adolescents, and seven adults. Paleobiology of the El Sidrón humans fits the pattern found in other neandertal samples: a high incidence of dental hypoplasia and interproximal grooves, yet no serious traumatic lesions are present. Moreover, unambiguous evidence of human-induced modifications (cannibalism) was found on the human remains: cut marks, percussion pitting, conchoidal scars and adhering flakes. Individuals seem to have been treated differentially. Morphologically, the El Sidrón humans show a large number of neandertal lineage-derived features even though certain traits place the sample at the limits of neandertal variation. Integrating the El Sidrón human mandibles and occipital bones into the larger neandertal sample reveals a possible geographic patterning, with southern Neandertals showing broader faces with increased lower facial heights. Ancient DNA analyses have been carried out, developing an anti-contamination protocol of excavation for minimizing the risk of modern human DNA contamination. As a result both mitochondrial and nuclear DNA have been extracted from dental and osteological remains. Curiously, mtDNA comparative analyses suggest a population affinity of Iberian Peninsula Neandertals with Central European Neandertals. Nuclear DNA analyses have permitted the identification of some functional genes such as the melanocortin 1 receptor (MC1R), which regulates hair and skin pigmentation; the FOXP2, a gene involved in the development of language; and the gene involved in the ABO blood group system. Nowadays the large El Sidrón sample is the most significant neandertal sample from the Iberian Peninsula, and augments the European evolutionary lineage fossil record, supporting ecogeographical variability across neandertal populations. [Copyright &y& Elsevier]

The distribution of subvertical grooves on interproximal wear dental facets from the El Sidrón (Asturias, Spain) Neandertals is described and analyzed. Out of 93 teeth, 64.5% present subvertical grooves, including a high frequency (50%) on the anterior dentition. Contrary to some studies, subvertical grooves from adjacent facets perfectly overlap each other and do not interdigitate, probably forming small channels. Both the facet and the groove surface share the same polished appearance, suggesting a common origin. Statistical analyses reveal that the number of grooves is neither dependent on the degree of occlusal wear, nor on the position on the tooth or the individual's age. However, facet width is an important factor determining the number of subvertical grooves. The etiology of subvertical grooves formation on Neandertal teeth remains unclear. Am [ABSTRACT FROM AUTHOR]

This paper presents the first detailed analysis of the lithic industry from El Sidrón Cave. Previously we have published other works concerning Anthropology and Palaeogenetics. The lithic assemblage contains 415 artefacts coming from both unofficial and official excavations and is largely flake-based. The raw material derives from the immediate cave environment, and is mostly chert, with quartzite in a lesser proportion. To date, 67 artefacts have been refitted, including some tools and cores. The spatial distribution of the refitted lithic artefacts confirms, on the one hand, the unity of the assemblage and, on the other, its secondary context. [ABSTRACT FROM AUTHOR]

Morphological differences between the Neandertal and modern human glenoid fossa are mostly attributed to functional factors. However, the intimate relationship between the pectoral girdle and the shape of the thorax on which it rests calls for a structural analysis of the morphology of the glenoid fossa. Using both 3D Geometric Morphometric techniques and classical osteometry we carry out an analysis of the morphology of Neandertal scapular glenoid fossa, considering the effects that independent parameters such as allometry, sexual dimorphism and laterality could exert on the glenoid fossa shape. We also discuss how the morphological configuration of the thorax together with other functional factors could explain some of the differences between Neandertals and H. sapiens , and we assess the morphological evolution of this structure in the context of the genus Homo . To this end, 3DGM methods were applied in a comparative framework including the new SD-2101 + specimen from the Sidrón site (Spain) dated to 49,000 years ago, other Neandertal, Homo sapiens , KNM-WT 15000 (Homo ergaster/erectus ) and Pan troglodytes specimens. A total of 75 sliding semilandmarks were used to collect the morphology of the glenoid fossa. Morphological comparison and variability were assessed through principal component analysis. Our results confirm the metric and morphological features of the Neandertal scapular glenoid fossa and point to a multifactorial effect in the glenoid morphology. We also detect a morphological trait of the Neandertal glenoid fossa that has not been reported previously: the projection of the upper part of the ventral border shared with modern humans and influenced by structural factors related to the configuration of the rib cage and shoulder. These results are consistent with the evolutionary framework of the hominin glenoid fossa, with some morphological changes being caused by functional factors, and others by structural factors such as the configuration of the thorax. [ABSTRACT FROM AUTHOR]

Abstract: This paper analyses the occipital remains recovered from the El Sidrón (Asturias, Spain) Neandertal site between the years of 2000–2008. The sample is represented by three specimens, SD-1219, SD-1149, and SD-370a. Descriptive morphology, linear measurements, 3D geometric morphometrics, and virtual anthropological methods were employed to address the morphological, morphometric, and phylogenetic affinities of these fossils. The fossils display Neandertal autapomorphies (e.g., bilaterally protruding transverse occipital torus, suprainiac fossa). SD-1219 also preserves a strongly projecting juxtamastoid eminence and shows occipital bunning. In linear distances, the El Sidrón occipitals are similar to each other and close to the Neandertal mean. The centroid size of SD-1219 is slightly larger than the Neandertal average. All of the evidence taken together points to the hypothesis that SD-1219 belongs to a smaller Neandertal male. Linear measurements and the vault thickness of SD-1149 also suggest a robust male individual. The gracility of SD-370a points towards an immature individual. Virtual anthropological methods were used to reconstruct a 3D model of the SD-1219 occipital for geometric morphometrics, which reveals that SD-1219 shows relatively broad and low occipital plane proportions. Within the European Pleistocene lineage sample, this fossil falls geometrically closer to primitive rather than to derived morphologies because of its increased width, and a lower, anterior position of inion relative to the biasterionic axis. These results may imply that cranial sphericity could be an important feature of intraspecific Neandertal variability. Our findings open the way for further studies of intraspecific variation in Neandertal populations, in which the El Sidrón sample may play a significant role. [Copyright &y& Elsevier]