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The BB specimen is an upper right canine belonging to a male of the primate M. sylvanus. The agreement among the qualitative characteristics of the tooth and the chronology of the site suggest its identification as M. sylvanus cf. florentina, although the lack of consensus on the validity of the proposed subspecies and the large biometric overlap do not allow certainty of this subspecific assignment. Macaca cf. sylvanus fossils are known across Iberia and throughout Europe in the Pliocene and Pleistocene, mainly in warmer intervals. Data suggest that the species is limited to areas that are more southern in colder phases, whereas in warm phases, this taxon could extend its distribution further north. Predation by carnivore species is likely, while interaction with early humans is less definite., This research has been funded by Ministerio de Ciencia e Innovacion, PID2021-122356NB-I00. D.F. is supported by the Ayuda del Programa de Formacion de Profesorado Universitario (FPU20/03389) and is a Ph.D. student at the Programa de Doctorado en Biología at the Universidad Complutense de Madrid. A.E. is supported by H2020-MSCA-IF project No. 891529 (3DFOSSILDIET). A.P. is supported by the LATEUROPE project (Grantagreement ID 101052653) that has received funding from the European Research Council (ERC) under the European Union’s HORIZON1.1 research program. The Barranc de la Boella fieldwork is supported by the Ajuntament de la Canonja and Departament de Cultura of Generalitat de Catalunya (ARQ001SOL-186-2022). The Institut Catala de Paleoecologia Humana i Evoluci o Social (IPHES- CERCA) has received financial support from the Spanish Ministry of Science and Innovation through the ‘María de Maeztu’ program for Units of Excellence (CEX2019-000945-M)., With funding from the Spanish government through the "Severo Ochoa Center of Excellence" accreditation CEX2019-000945-M.

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. We thank the curators at the following museums for providing access to the collections: Museo de Arqueología de Asturias (Las Caldas and Tito Bustillo), Museo de Prehistoria y Arqueología de Cantabria (El Castillo) and Arkeologi Museoa (Santa Catalina and Axlor).We are very grateful to the Editors-in-Chief (David Alba and Clément Zanolli), the Associate Editor, and reviewers for their comments and suggestions, which greatly improved the quality of this paper. A.E. was beneficiary of a Juan de la Cierva-Formación Postdoctoral Fellowship (grant number FJCI-2016-30122). Partial aspects of this research have been founded by SUBSILIENCE ERCProject (ERCEA-; grant agreement No. 818299), ABRUPT project (HAR2017-84997-P Spanish Ministry of Science, Innovation and Universities) and Santander Bank within the program of Santander Talent Attraction for Research (STAR1) to A.B.M.A., and H2020- MSCA-IF project No. 891529 (3DFOSSILDIET) to A.E. This project has received funding from the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement No 891529

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., This work is funded by the Spanish Ministry of Economy and Competitiveness: CGL2016-75109-P.

Objectives The El Sidron 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.

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., This research was funded by the local Government of the Principado de Asturias and by the Spanish Government (Project CGL2012-36682), and the European Community Research Infrastructure Action (SYNTHESYS Project; http://www.synthesys.info, to S. El Zaatari).

Objectives During the microscopic examination of the Neandertal dentitions from El Sidron (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 Sidron individuals (nine out of 11). Toothpick grooves from El Sidron 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.

[EN] 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., [ES] El estudio del tórax Neandertal ha atraído el interés de la comunidad científica por más de un siglo. Existe acuerdo acerca de la mayor capacidad torácica en Neandertales en comparación con humanos modernos. Sin embargo, si esto es causado por una expansión antero-posterior o medio-lateral del tórax es debatido a día de hoy y es clave para el entendimiento de la biomecánica respiratoria y la morfología corporal en Neandertales. La frágil naturaleza de las costillas, la estructura metamérica del tórax y las dificultades en la cuantificación morfológica contribuyen a la incertidumbre en referencia a aspectos de la morfología torácica Neandertal. El yacimiento de El Sidrón ha proporcionado elementos costales que comprenden desde el tórax superior hasta el inferior, así como diferentes restos costales proximales (frecuentemente ausentes en el registro fósil Neandertal), los cuales pueden arrojar luz sobre esta incertidumbre. Nosotros comparamos las costillas de El Sidrón con costillas de humanos modernos actuales, así como con humanos modernos fósiles y otros Neandertales, a través de técnicas de morfometría clásica y morfometría geométrica 3D, combinadas con técnicas de estimación de datos perdidos y de reconstrucción virtual (a nivel de 1a, 5a y 11a costilla). Nuestros resultados muestran que los Neandertales presentan cabezas costales y tubérculos articulares más grandes que humanos modernos. A nivel de tamaño global de las costillas, las 1a costillas Neandertales son más peque~nas que las de humanos modernos, mientras que las 5a y 11a son considerablemente más grandes. Cuando articulamos costillas medias (forma y tamaño) con sus correspondientes elementos vertebrales, nosotros observamos que el tórax Neandertal presenta una expansión medio-lateral en los diferentes niveles estudiados con respecto a humanos modernos, aunque esto es más evidente a nivel T5 y T11. Por lo tanto, a la luz de la evidencia proporcionada por los restos costales de El Sidrón, nosotros hipotetizamos que la expansión volumétrica Neandertal propuesta por autores previos, debería ser fundamentalmente producida por una expansión medio-lateral del tórax., This study was funded by The Leakey Foundation, the Ministerio de Economía y Competitividad of Spain (CGL2012-36682, CGL2015-63648-P and CGL2012-37279) and the Convenio Principado de Asturias-Universidad de Oviedo (CN-09-084). MB and DGM were funded by the European Commission's Research Infrastructure Action via the Synthesys Projects (AT-TAF-4616 for MB and DK-TAF-3494 and BE-TAF-5639 for DGM).

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., This research was supported by the European Research Council Starting Grant 336301 and the Max Planck Society (TLK), and the Ministerio de Economía y Competitividad of Spain CGL2016-75109-P.

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., Portions of this work was funded by the Dirección General de Investigación of MINECO, projects number CGL2012‐36682, CGL2015‐65387‐C3‐1‐P, CGL2015‐65387‐C3‐2‐P, CGL2015‐65387‐C3‐3‐P (MINECO/FEDER), CERCA Programme and project AGAUR 2014SGR900 of Generalitat de Catalunya.

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.

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., This work was funded by the Ministerio de Economía y Competitividad of Spain (CGL2012-36682 and CGL2012-37279) and the Convenio Principado de Asturias-Universidad de Oviedo (CN-09-084).

Well preserved thoracic vertebrae of Neandertals are rare. However, such fossils are im portant as t heir three-dimensio nal (3D) spatial con ¿ guration can contr ibute to the under standing of the size and shape of the thoracic spine and the entire thorax. This is be cause the verteb ral body and t ransverse processes provid e the articulati on and attachme nt si tes for the ribs . Dorsal ori entation of the transverse processes relative to the vertebral body also rotates the attache d ribs in a way that could affe ct thorax wi dth. Previous research indicates pos sible evide nce for greater dorsal orien tation of the transverse processes and small vertebral body heig hts in Neandertals, but their 3D verteb ral str ucture has not yet be en addressed. Here we present 15 new vertebral remains from the El Si dr on Neandertals (Ast urias, Northern Spain ) and used 3D geometric morph ometri cs to address the above issues by comp aring two par ticularly well preserved El Sidr on remains (SD-1619, SD-16 41) with thoracic vertebrae from ot her Neandertals and a sa mple of anatomically mod ern human s. Centroid size s of El Si dron vertebrae are withi n the human range. Neandertals have larger T1 and probably also T2. The El Sidron vertebrae are similar in 3D shap e to those of other Neandertals, wh ich differ from Homo sapien s par ticularly in central-lower regions (T6 e T10) of the thoracic spine. Differences in clude more dorsal ly and cranially oriented t ransverse pro-cesse s, less cau dally ori ented spino us processes, and vertebral bodies that are anteroposter iorly and craniocaudally short. The results ¿ t wit h current reconstruc tions of Neandertal t horax morphology, This research is funded by CGL2012-37279, CGL2012-36682, and CGL2015-63648-P (Ministerio de Economía y Competitividad, Spain)

Received 25 October 2013, Accepted 8 July 2014, Available online 11 February 2015, [EN] 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., [ES] El patrón de desgaste dental no masticatorio de los cazadores-recolectores prehistóricos y grupos actuales, muestra diferencias según del sexo de los individuos, reflejando una división sexual del trabajo. Los Neandertales, por su parte, tienen un gran desgaste de sus incisivos y caninos debido a que los usarían como coadyuvante o ‘tercera mano’ en numerosas tareas, como la preparación de pieles o el troceado de pedazos de carne que sujetaban con la boca. En este artículo se presenta el estudio del desgaste dental no masticatorio en 19 individuos neandertales de los yacimientos de l'Hortus (Francia), Spy (Bélgica) y El Sidrón (España). Para ello, se han analizado dos de los rasgos asociados a este tipo de desgaste dental: las estrías culturales (rayas superficiales hechas en la cara labial de la dentición anterior y que tienen la misma morfología que una marca de corte) y las melladuras del esmalte dental (fracturas producidas ante-mortem que afectan al esmalte dental o bien al esmalte y a la dentina en conjunto). Los resultados muestran que todos los individuos presentan ambos rasgos de desgaste no-masticatorio, pero se han encontrado diferencias significativas en cuanto a la longitud de las estrías, siendo más largas en las mujeres. En cuanto a la distribución de las mellas en el esmalte, los hombres tienen una mayor incidencia en la dentición maxilar, mientras que las mujeres es la mandibular. Todas las diferencias detectadas en el patrón de desgaste dental no masticatorio, aunque sutiles, parecen indicar que los Neandertales utilizaban sus dientes de manera diferente según el sexo, sugiriendo que un cierto grado de división del trabajo ya estaba presente en esta especie humana extinta., A. E. is the beneficiary of a predoctoral research grant (FPI; BES-2010-033914)., This research has been funded by the local Government of the Principado de Asturias and the Spanish Government (Projects CGL2012-36682).

© 2014 Elsevier Ltd. 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, This project is funded by the Leakey Foundation, and by the Spanish Ministry of Economy and Competitivity: CGL2012-37279 (MB), and CGL2012-36682 (AR).

Received 18 March 2014, Accepted 27 February 2015, Available online 24 March 2015, 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., Ministerio de Economía y Competitividad of Spain: CGL2012-36682 and CGL2012-37279., Convenio Principado de Asturias-Universidad de Oviedo CN-09-084.

Josep Vallverdú [et al.].- Received March 2, 2014; Accepted June 30, 2014; Published July 30, 2014, 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. © 2014 Vallverdú et al., The research at Barranc de la Boella has been carried out with the financial support of the Spanish Ministerio de Economía y Competitividad (CGL2012-36682; CGL2012-38358, CGL2012-38434-C03-03 and CGL2010-15326; MICINN project HAR2009-7223/HIST), Generalitat de Catalunya, AGAUR agence (projects 2014SGR-901; 2014SGR-899; 2009SGR-324, 2009PBR-0033 and 2009SGR-188) and Junta de Castilla y León BU1004A09.

Received: February 28, 2013; Accepted: March 25, 2013; Published: May 6, 2013, 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., This research is funded by the Spanish Ministerio de Economía y Competitividad (Projects CGL2009-09013 and CGL2012-36682)., A.E. has the benefit of a predoctoral FPI fellowship of the Spanish Ministerio de Economía y Competitividad (BES-2010-033914) (http://www.mineco.gob.es/portal/site/mineco/).

Two Neandertal specimens from El Sidrón, northern Spain, show evidence of retained left mandibular deciduous canines. These individuals share the same mitochondrial (mtDNA) haplotype, indicating they are maternally related and suggesting a potential heritable basis for these dental anomalies. Radiographs and medical CT scans provide evidence of further, more extensive dental pathology in one of these specimens. An anomalous deciduous canine crown morphology that developed before birth subsequently suffered a fracture of the crown exposing the pulp sometime after eruption into functional occlusion. This led to death of the tooth, periapical granuloma formation and arrested deciduous canine root growth at an estimated age of 2.5 years. At some point the underlying permanent canine tooth became horizontally displaced and came to lie low in the trabecular bone of the mandibular corpus. A dentigerous cyst then developed around the crown. Anterior growth displacement of the mandible continued around the stationary permanent canine, leaving it posteriorly positioned in the mandibular corpus by the end of the growth period beneath the third permanent molar roots, which, in turn, suggests a largely horizontal growth vector. Subsequent longstanding repeated infections of the expanding cyst cavity are evidenced by bouts of bone deposition and resorption of the boundary walls of the cyst cavity. This resulted in the establishment of two permanent bony drainage sinuses, one through the buccal plate of the alveolar bone anteriorly, immediately beneath the infected deciduous canine root, and the other through the buccal plate anterior to the mesial root of the first permanent molar. It is probable that this complicated temporal sequence of dental pathologies had an initial heritable trigger that progressed in an unusually complex way in one of these individuals. During life, this individual may have been largely unaware of this ongoing pathology. © 2013 Elsevier Ltd., El Sidrón site is being excavated with the support of the Consejería de Cultura of the Asturias Autonomous government. This research is founded by the Spanish Ministry of Economy and Competitiveness (Project CGL2012-36682).

The distribution of subvertical grooves on interproximal wear dental facets from the El Sidrõn (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. © 2010 Wiley-Liss, Inc., Funded by: Universidad de Oviedo-CSIC. Grant Number: 060501040023, Ministerio de Ciencia e Innovación of Spain. Grant Number: CGL2009-09013