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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]

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. [ABSTRACT FROM AUTHOR]

Competing Interests: Declaration of competing interest The authors of this manuscript certify that they have no conflict of interest. They have no affiliations with or involvement in any organization or entity with any financial interest (such as honoraria; educational grants; participation in speakers’ bureaus; membership, employment, consultancies, stock ownership, or other equity interest; and expert testimony or patent-licensing arrangements), or non-financial interest (such as personal or professional relationships, affiliations, knowledge or beliefs) in the subject matter or materials discussed in this manuscript.

Objective: To encourage the use of different methodological approaches for the identification of paleopathological lesions and to evaluate osteolytic lesions found on a temporal bone from La Llana cave (Spain)., Materials: Cranial remains recovered from from La Llana cave (Spain) dated to the Bronze Age, 3300 ± 25 BP (1631-1509 cal BC)., Methods: The cranium underwent macroscopic, microscopic and computed tomography scan examinations., Results: The Tegmen tympani of the left temporal bone is present but is missing on the right. Both cochleae are intact. Both the right and left temporal bones display osteolytic lesions, with the left Tegmen tympani displaying deep and profuse pits on the endocranial surface, exposing the ear channel., Conclusions: The lesions are compatible with otitis media (atticitis). By using different methods of analysis, the presence and effects of infection were identified., Significance: This systematically described case contributes to our understanding of disease in the past and describes otitis-related lesions in archeological contexts for comparative purposes., Limitations: The fragmentary and damaged cranial remains rendered evaluation difficult and diagnosis tentative., Suggestions for Further Research: The use of different approaches to diagnose otitis media in archeological skeletal collections is recommended to improve the knowledge of health status and lifestyle of past populations., (Copyright © 2020 Elsevier Inc. 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.)

We assembled genome-wide data from 271 ancient Iberians, of whom 176 are from the largely unsampled period after 2000 BCE, thereby providing a high-resolution time transect of the Iberian Peninsula. We document high genetic substructure between northwestern and southeastern hunter-gatherers before the spread of farming. We reveal sporadic contacts between Iberia and North Africa by ~2500 BCE and, by ~2000 BCE, the replacement of 40% of Iberia's ancestry and nearly 100% of its Y-chromosomes by people with Steppe ancestry. We show that, in the Iron Age, Steppe ancestry had spread not only into Indo-European-speaking regions but also into non-Indo-European-speaking ones, and we reveal that present-day Basques are best described as a typical Iron Age population without the admixture events that later affected the rest of Iberia. Additionally, we document how, beginning at least in the Roman period, the ancestry of the peninsula was transformed by gene flow from North Africa and the eastern Mediterranean., (Copyright © 2019 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Neandertals disappeared from the fossil record around 40,000 bp, after a demographic history of small and isolated groups with high but variable levels of inbreeding, and episodes of interbreeding with other Paleolithic hominins. It is reasonable to expect that high levels of endogamy could be expressed in the skeleton of at least some Neandertal groups. Genetic studies indicate that the 13 individuals from the site of El Sidrón, Spain, dated around 49,000 bp, constituted a closely related kin group, making these Neandertals an appropriate case study for the observation of skeletal signs of inbreeding. We present the complete study of the 1674 identified skeletal specimens from El Sidrón. Altogether, 17 congenital anomalies were observed (narrowing of the internal nasal fossa, retained deciduous canine, clefts of the first cervical vertebra, unilateral hypoplasia of the second cervical vertebra, clefting of the twelfth thoracic vertebra, diminutive thoracic or lumbar rib, os centrale carpi and bipartite scaphoid, tripartite patella, left foot anomaly and cuboid-navicular coalition), with at least four individuals presenting congenital conditions (clefts of the first cervical vertebra). At 49,000 years ago, the Neandertals from El Sidrón, with genetic and skeletal evidence of inbreeding, could be representative of the beginning of the demographic collapse of this hominin phenotype.

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.)

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 first arrivals of hominin populations into Eurasia during the Early Pleistocene are currently considered to have occurred as short and poorly dated biological dispersions. Questions as to the tempo and mode of these early prehistoric settlements have given rise to debates concerning the taxonomic significance of the lithic assemblages, as trace fossils, and the geographical distribution of the technological traditions found in the Lower Palaeolithic record. Here, we report on the Barranc de la Boella site which has yielded a lithic assemblage dating to ∼1 million years ago that includes large cutting tools (LCT). We argue that distinct technological traditions coexisted in the Iberian archaeological repertoires of the late Early Pleistocene age in a similar way to the earliest sub-Saharan African artefact assemblages. These differences between stone tool assemblages may be attributed to the different chronologies of hominin dispersal events. The archaeological record of Barranc de la Boella completes the geographical distribution of LCT assemblages across southern Eurasia during the EMPT (Early-Middle Pleistocene Transition, circa 942 to 641 kyr). Up to now, chronology of the earliest European LCT assemblages is based on the abundant Palaeolithic record found in terrace river sequences which have been dated to the end of the EMPT and later. However, the findings at Barranc de la Boella suggest that early LCT lithic assemblages appeared in the SW of Europe during earlier hominin dispersal episodes before the definitive colonization of temperate Eurasia took place.

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)

The remains of 12 Neandertal individuals have been found at the El Sidrón site (Asturias, Spain), consisting of six adults, three adolescents, two juveniles, and one infant. Archaeological, paleontological, and geological evidence indicates that these individuals represent all or part of a contemporaneous social group of Neandertals, who died at around the same time and later were buried together as a result of a collapse of an underground karst. We sequenced phylogenetically informative positions of mtDNA hypervariable regions 1 and 2 from each of the remains. Our results show that the 12 individuals stem from three different maternal lineages, accounting for seven, four, and one individual(s), respectively. Using a Y-chromosome assay to confirm the morphological determination of sex for each individual, we found that, although the three adult males carried the same mtDNA lineage, each of the three adult females carried different mtDNA lineages. These findings provide evidence to indicate that Neandertal groups not only were small and characterized by low genetic diversity but also were likely to have practiced patrilocal mating behavior.

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.