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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.

At the unique site of El Sidrón there have been an impressive series of events related to the activities and behaviors of the Neandertal species, integrated into peculiar environmental processes, seasoned with the proximity of the best Asturian siliceous raw material, and with a peculiar chain of expeditious and fast situations. All this integrated into an interdisciplinary investigation, En el singular yacimiento de El Sidrón se han producido una impactante serie de acontecimientos relacionados con las actividades y comportamientos de la especie neandertal, integradas en unos procesos del medio físico peculiares, sazonadas con la proximidad de la mejor materia prima silícea asturiana, y con un peculiar encadenamiento de situaciones expeditivas y rápidas. Todo ello integrado en una investigación interdisciplinar

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 geneti

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.

This study presents the results of the archaeological investigation in Benzú Cave, located on the North African shore of the Strait of Gibraltar. The archaeological deposits, approximately 1 m deep, belong to two occupational levels dated to the Neolithic and the Bronze Age. A wide range of artifacts was found, and this led to an interdisciplinary study that led us to reach new conclusions concerning the material life, subsistence economy, and environment of the Neolithic and Bronze Age societies in northern Africa and the southern Iberian Peninsula.

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 thor

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.

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.

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.

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.

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

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.

The late Early Pleistocene site Barranc de la Boella provides an unparalleled opportunity to assess the context of the activities of the hominin populations that inhabited the Iberian Peninsula at 1 Ma. Recently, strong evidence for access to mammoth meat has been described at the Pit 1 locality. At the la Mina and el Forn excavation areas, little evidence exists for the anthropogenic processing of macromammals. However, the presence of humans is recorded, and the available evidence suggests these populations had access to several interesting resources. By analysing and comparing ¿ve separate assemblages at Barranc de la Boella, we assess the ¿uctuating presence of hominins and carnivores and the levels of competition among predators in each assemblage. Our analysis re-veals different levels of competition intensity during the formation of assemblages when hominin groups were present, as evidenced by the abundance and diversity of stone artefacts. The analyses of skeletal component ratios indicate several competitive contexts, and the greatest presence of hominin groups is associated with the most c om pe ti t i ve sc e n a ri os . T h e p al a e oe n vi r on m e nt a t Ba r ra nc de l a Bo e ll a wa s r i ch i n r e so u rc e s t ha t ho m in i n s could exploit. The presence of hominin and carnivore groups appears to have been higher in levels with more in-ferred competition. This scenario supports prior research that concludes that carnivore abundance and highly competitive contexts were two constants in the lives of these hominin groups. Thus, the criteria determining wh e th e r ho m i n i ns c ou l d i nh a bi t a gi v e n l a nd s ca pe we re m os t l i k e l y r el a te d to th e pr e se n ce or ab se n ce of resources, such as animal resources, water and raw materials, rather than the dynamics of the carnivore popula-tions, to which the hominins were able to become habituated.

Coprolites are commonly identified in the Pleistocene archaeo-palaeontological record. They have often been described as indirect evidence for the presence of carnivores (usually hyenids) during the formation of a depositional sequence. However, coprolites are a much larger source of information that can provide data relating to factors affecting an archaeological assemblage, including its taphonomic history and palaeoecology. In this paper, two fossil coprolite accumulations dating to the late Early Pleistocene are described: Level TD6.1 of Gran Dolina (Sierra de Atapuerca, Burgos) and Unit II of la Mina (Barranc de la Boella, Tarragona). The results highlight the morphological homogeneity of these hyena coprolites, despite a considerable variation in size. The presence of microspherulites is clearly identified in all of the analysed hyena coprolites. Their composition is rich in elements characteristic of bone-origin, such as calcium (Ca) and phosphorus (P). A major difference, however, differentiating the remains from these sites was a higher frequency of bone fragments in the la Mina coprolites, regarding to TD6.1. Pollen, spores, and palynomorphs are scarce in the samples from both sites, which in fact share similar ecological characteristics such as the presence of pine, wild grasses, and coprophilous fungi. Hyena coprolites are relatively easy to differentiate from those of other taxa; however, they show intra-specific similarities in their micro- and macro-morphology and composition, despite their wide size range. The species of hyenids responsible for the coprolites is not possible to identify based exclusively on the size range, and the importance of a multidisciplinary study of the latrines and the coprolites is discussed as a source of palaeoecological and taphonomic information. In this study, coprolites are analysed, compared, and attributed to a hyenid, identified as Crocuta crocuta in the case of TD6.1, but without specific attribution in the case of

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.

[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 h

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 t o b e 7.7 years. Maturation of most elements fell within the expected range of modern humans at this age. The exce ptions were the atlas and mid-thoraci c ver tebrae, whi ch remai ned a t t he 5- to 6-ye a r sta ge o f d evelopment. Furthermo re, e ndocra nial features suggestthatbraingrowth wasnot yetcompleted. T he vertebral m aturation p attern 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.