Death in the high mountains: Evidence of interpersonal violence during Late Chalcolithic and Early Bronze Age at Roc de les Orenetes (Eastern Pyrenees, Spain).

OBJECTIVES: To test a hypothesis on interpersonal violence events during the transition between Chalcolithic and Bronze Age in the Eastern Pyrenees, to contextualize it in Western Europe during that period, and to assess if these marks can be differentiated from secondary funerary treatment. MATERIALS AND METHODS: Metric and non-metric methods were used to estimate the age-at-death and sex of the skeletal remains. Perimortem injuries were observed and analyzed with stereomicroscopy and confocal microscopy. RESULTS: Among the minimum of 51 individuals documented, at least six people showed evidence of perimortem trauma. All age groups and both sexes are represented in the skeletal sample, but those with violent injuries are predominantly males. Twenty-six bones had 49 injuries, 48 of which involved sharp force trauma on postcranial elements, and one example of blunt force trauma on a cranium. The wounds were mostly located on the upper extremities and ribs, anterior and posterior. Several antemortem lesions were also documented in the assemblage. DISCUSSION: The perimortem lesions, together with direct dating, suggest that more than one episode of interpersonal violence took place between the Late Chalcolithic and the Early Bronze Age in northeastern Spain. The features of the sharp force trauma indicate that different weapons were used, including sharp metal objects and lithic projectiles. The Roc de les Orenetes assemblage represents a scenario of recurrent lethal confrontation in a high mountain geographic context, representing the evidence of inferred interpersonal violence located at the highest altitude settings in the Pyrenees, at 1836 meters above sea level.

OBJETIVOS: Estudiar nuevas evidencias de violencia interpersonal durante la transición entre el Calcolítico y la Edad del Bronce en los Pirineos Orientales, contextualizarlas en la Europa occidental durante ese periodo, y diferenciar estas marcas del tratamiento funerario secundario. MATERIALES Y MÉTODOS: Se han utilizado métodos métricos y no métricos para estimar la edad de muerte y sexo de los restos esqueléticos. Las heridas perimortem fueron observadas y analizadas con estereomicroscopio y microscopio confocal. RESULTADOS: Entre el mínimo de 51 individuos documentados en el yacimiento, al menos seis individuos mostraron evidencias de traumas perimortem. Todos los grupos de edad y ambos sexos están representados en el enterramiento, pero aquellos con heridas violentas son mayoritariamente masculinos. Veintiséis huesos tenían un total de 49 lesiones, 48 de ellos traumatismos cortantes en elementos postcraneales, y un traumatismo contundente en un cráneo. Las heridas estaban mayoritariamente localizadas en las extremidades superiores y costillas, tanto anterior como posterior. Varias lesiones antemortem fueron también documentadas en el conjunto. DISCUSIÓN: Las lesiones perimortem, junto a las dataciones directas, sugieren que se produjo más de un episodio de violencia interpersonal entre el Calcolítico Final y la Edad del Bronce Antiguo en el noreste de España. Las características de los traumatismos cortantes indican que se utilizaron diferentes armas, incluyendo objetos cortantes de metal y proyectiles líticos. El conjunto de Roc de les Orenetes representa un escenario de confrontaciones letales recurrentes en un contexto geográfico de alta montaña, representando la evidencia de violencia interpersonal localizada a mayor altitud de los Pirineos, a 1836 metros sobre el nivel del mar.

Unearthing Neanderthal population history using nuclear and mitochondrial DNA from cave sediments.

Bones and teeth are important sources of Pleistocene hominin DNA, but are rarely recovered at archaeological sites. Mitochondrial DNA (mtDNA) has been retrieved from cave sediments but provides limited value for studying population relationships. We therefore developed methods for the enrichment and analysis of nuclear DNA from sediments and applied them to cave deposits in western Europe and southern Siberia dated to between 200,000 and 50,000 years ago. We detected a population replacement in northern Spain about 100,000 years ago, which was accompanied by a turnover of mtDNA. We also identified two radiation events in Neanderthal history during the early part of the Late Pleistocene. Our work lays the ground for studying the population history of ancient hominins from trace amounts of nuclear DNA in sediments.

Neanderthals and Homo sapiens had similar auditory and speech capacities.

The study of audition in fossil hominins is of great interest given its relationship with intraspecific vocal communication. While the auditory capacities have been studied in early hominins and in the Middle Pleistocene Sima de los Huesos hominins, less is known about the hearing abilities of the Neanderthals. Here, we provide a detailed approach to their auditory capacities. Relying on computerized tomography scans and a comprehensive model from the field of auditory bioengineering, we have established sound power transmission through the outer and middle ear and calculated the occupied bandwidth in Neanderthals. The occupied bandwidth is directly related to the efficiency of the vocal communication system of a species. Our results show that the occupied bandwidth of Neanderthals was greater than the Sima de los Huesos hominins and similar to extant humans, implying that Neanderthals evolved the auditory capacities to support a vocal communication system as efficient as modern human speech.

Author Correction: The dental proteome of Homo antecessor.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

The dawn of the Middle Paleolithic in Atapuerca: the lithic assemblage of TD10.1 from Gran Dolina.

The Atapuerca localities present evidence of a long series of hominin occupations from the Early Pleistocene onward and are a key site for understanding the continuity and discontinuity of Western European technological and settlement dynamics. The TD10 unit from Gran Dolina is located in the upper part of the sequence and divided into four lithostratigraphic subunits (TD10.4 to TD10.1, from bottom to top) dated between ca. 450 ka and ca. 250 ka (Marine Isotope Stage 11 to Marine Isotope Stage 8). The technological analysis of the lithic assemblages belonging to the TD10.1 sequence aims to determine the trends among its archeological levels and check its relation to late Middle Pleistocene technological evolution and site functionality. Archeostratigraphic studies have identified several occupation events within its approximately 1.5 m of thickness, whose artifact densities and occupational models differ. However, no remarkable technical differences have been observed among them. Lithic assemblages from those events show more evolved features than other Atapuerca Mode 2 assemblages. These changes are reflected in the selective raw material management strategies; more hierarchized and predetermined reduction methods; and the progressive decrease of large cutting tools in the lithic assemblages with respect to flake tools, the latter defined by a greater typological diversification. These technological changes did not lead to a clear break with respect to previous technological models and were accompanied by other sporadic but significant changes in subsistence and behavioral strategies (bone tools and retouchers; lithic recycling, and so on), which were consolidated during the Middle Paleolithic. Hence, the archeological record from the TD10.1 subunit of Gran Dolina reflects a local stratigraphic transition from Mode 2 to Mode 3 technocomplexes, paralleling that observed in other sites in southwestern Europe.

The dental proteome of Homo antecessor.

The phylogenetic relationships between hominins of the Early Pleistocene epoch in Eurasia, such as Homo antecessor, and hominins that appear later in the fossil record during the Middle Pleistocene epoch, such as Homo sapiens, are highly debated1-5. For the oldest remains, the molecular study of these relationships is hindered by the degradation of ancient DNA. However, recent research has demonstrated that the analysis of ancient proteins can address this challenge6-8. Here we present the dental enamel proteomes of H. antecessor from Atapuerca (Spain)9,10 and Homo erectus from Dmanisi (Georgia)1, two key fossil assemblages that have a central role in models of Pleistocene hominin morphology, dispersal and divergence. We provide evidence that H. antecessor is a close sister lineage to subsequent Middle and Late Pleistocene hominins, including modern humans, Neanderthals and Denisovans. This placement implies that the modern-like face of H. antecessor-that is, similar to that of modern humans-may have a considerably deep ancestry in the genus Homo, and that the cranial morphology of Neanderthals represents a derived form. By recovering AMELY-specific peptide sequences, we also conclude that the H. antecessor molar fragment from Atapuerca that we analysed belonged to a male individual. Finally, these H. antecessor and H. erectus fossils preserve evidence of enamel proteome phosphorylation and proteolytic digestion that occurred in vivo during tooth formation. Our results provide important insights into the evolutionary relationships between H. antecessor and other hominin groups, and pave the way for future studies using enamel proteomes to investigate hominin biology across the existence of the genus Homo.

Short and long period growth markers of enamel formation distinguish European Pleistocene hominins.

Characterizing dental development in fossil hominins is important for distinguishing between them and for establishing where and when the slow overall growth and development of modern humans appeared. Dental development of australopiths and early Homo was faster than modern humans. The Atapuerca fossils (Spain) fill a barely known gap in human evolution, spanning ~1.2 to ~0.4 million years (Ma), during which H. sapiens and Neandertal dental growth characteristics may have developed. We report here perikymata counts, perikymata distributions and periodicities of all teeth belonging to the TE9 level of Sima del Elefante, level TD6.2 of Gran Dolina (H. antecessor) and Sima de los Huesos. We found some components of dental growth in the Atapuerca fossils resembled more recent H. sapiens. Mosaic evolution of perikymata counts and distribution generate three distinct clusters: H. antecessor, Sima de los Huesos and H. sapiens.

Ectopic maxillary third molar in Early Pleistocene Homo antecessor from Atapuerca-Gran Dolina site (Burgos, Spain).

OBJECTIVES: Here we describe the case of an ectopic maxillary third molar (M3 ), preventing the eruption of the M2 , in the individual H3 of the hominin hypodigm of level TD6.2 of the Early Pleistocene site of Gran Dolina (Sierra de Atapuerca, Spain). MATERIALS AND METHODS: The fossil remains from the TD6.2 level of the Gran Dolina site (about 170 specimens) are assigned to Homo antecessor. Different geochronological methods place these hominins in the oxygen isotopic stage 21, between 0.8 and 0.85 million years ago (Ma). The immature individual H3 is represented by an almost complete midface (ATD6-69), preserving various teeth in situ. We used high-resolution microtomograhy (mCT) to investigate the abnormal position of the left M3 , virtually reconstruct M2 , and M3 as well as assessing the development stage of these. Finally, we compare this case with extinct and extant populations. RESULTS: Based on the identified signs, we suggest that individual H3 suffered from a unilateral impaction of the M2 as a result of the ectopic position of the developing M3 . DISCUSSION: We conclude that the most likely etiology for the ectopic position of the M3 is the lack of space in the maxilla. We discuss possible contributing factors, such as morphometric aspects of the maxilla and the early mineralization of the M3 , to support the M2 impaction. Finally, due to the early age at death of this individual we did not identify any secondary lesion associated with the M2 impaction.

Four millennia of Iberian biomolecular prehistory illustrate the impact of prehistoric migrations at the far end of Eurasia.

Population genomic studies of ancient human remains have shown how modern-day European population structure has been shaped by a number of prehistoric migrations. The Neolithization of Europe has been associated with large-scale migrations from Anatolia, which was followed by migrations of herders from the Pontic steppe at the onset of the Bronze Age. Southwestern Europe was one of the last parts of the continent reached by these migrations, and modern-day populations from this region show intriguing similarities to the initial Neolithic migrants. Partly due to climatic conditions that are unfavorable for DNA preservation, regional studies on the Mediterranean remain challenging. Here, we present genome-wide sequence data from 13 individuals combined with stable isotope analysis from the north and south of Iberia covering a four-millennial temporal transect (7,500-3,500 BP). Early Iberian farmers and Early Central European farmers exhibit significant genetic differences, suggesting two independent fronts of the Neolithic expansion. The first Neolithic migrants that arrived in Iberia had low levels of genetic diversity, potentially reflecting a small number of individuals; this diversity gradually increased over time from mixing with local hunter-gatherers and potential population expansion. The impact of post-Neolithic migrations on Iberia was much smaller than for the rest of the continent, showing little external influence from the Neolithic to the Bronze Age. Paleodietary reconstruction shows that these populations have a remarkable degree of dietary homogeneity across space and time, suggesting a strong reliance on terrestrial food resources despite changing culture and genetic make-up.

Shedding light on the Early Pleistocene of TD6 (Gran Dolina, Atapuerca, Spain): The technological sequence and occupational inferences.

This paper aims to update the information available on the lithic assemblage from the entire sequence of TD6 now that the most recent excavations have been completed, and to explore possible changes in both occupational patterns and technological strategies evidenced in the unit. This is the first study to analyse the entire TD6 sequence, including subunits TD6.3 and TD6.1, which have never been studied, along with the better-known TD6.2 Homo antecessor-bearing subunit. We also present an analysis of several lithic refits found in TD6, as well as certain technical features that may help characterise the hominin occupations. The archaeo-palaeontological record from TD6 consists of 9,452 faunal remains, 443 coprolites, 1,046 lithic pieces, 170 hominin remains and 91 Celtis seeds. The characteristics of this record seem to indicate two main stages of occupation. In the oldest subunit, TD6.3, the lithic assemblage points to the light and limited hominin occupation of the cave, which does, however, grow over the course of the level. In contrast, the lithic assemblages from TD6.2 and TD6.1 are rich and varied, which may reflect Gran Dolina cave's establishment as a landmark in the region. Despite the occupational differences between the lowermost subunit and the rest of the deposit, technologically the TD6 lithic assemblage is extremely homogeneous throughout. In addition, the composition and spatial distribution of the 12 groups of lithic refits found in unit TD6, as well as the in situ nature of the assemblage demonstrate the high degree of preservation at the site. This may help clarify the nature of the Early Pleistocene hominin occupations of TD6, and raise reasonable doubt about the latest interpretations that support the ex situ character of the assemblage as a whole.

Right-handed fossil humans.

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.

Neanderthal hunting strategies inferred from mortality profiles within the Abric Romaní sequence.

Ungulate mortality profiles are commonly used to study Neanderthal subsistence strategies. To assess the hunting strategies used by Neanderthals, we studied the ages at death of the cervids and equids found in levels E, H, I, Ja, Jb, K, L and M of the Abric Romaní sequence. These levels date between 43.2 ± 1.1 ka BP (14C AMS) and 54.5 ± 1.7 ka BP (U-series). The degree of eruption and development of the teeth and their wear stages were used to determine the ages of these animals at death, and mortality profiles were constructed using these data. The equids display prime dominated profiles in all of the analyzed levels, whereas the cervids display variable profiles. These results suggest that the Neanderthals of Abric Romaní employed both selective and non-selective hunting strategies. The selective strategy focused on the hunting of prime adults and generated prime dominated profiles. On the other hand, non-selective strategies, involved the consumption of animals of variable ages, resulting in catastrophic profiles. It is likely that in the selective hunting events were conducted using selective ambushes in which it was possible to select specific prey animals. On the other hand, encounter hunting or non-selective ambush hunting may have also been used at times, based on the abundances of prey animals and encounter rates. Specific hunting strategies would have been developed accordance with the taxa and the age of the individual to be hunted. The hunting groups most likely employed cooperative hunting techniques, especially in the capture of large animals. Thus, it is not possible to uniquely associate a single mortality profile with the predation tactics of Neanderthals at Abric Romaní.

Twentieth anniversary of Homo antecessor (1997-2017): a review.

It has been twenty years since diagnosis and publication of the species Homo antecessor.1 Since then, new human fossils recovered from the TD6 level of the Gran Dolina site (Sierra de Atapuerca, northern Spain) have helped to refine its taxonomic and phylogenetic position. In this paper, we present a synthesis of the most characteristic features of this species, as well as our interpretation derived from the latest investigations. We focus on the phylogenetic interpretation of Homo antecessor, taking into account the most recent paleogenetic analyses and a reassessment of the European Middle Pleistocene hominin record. We try to show that, twenty years after its publication, H. antecessor provides a good opportunity to address the morphology of the last common ancestor of Neandertals and modern humans.

Human predatory behavior and the social implications of communal hunting based on evidence from the TD10.2 bison bone bed at Gran Dolina (Atapuerca, Spain).

Zooarcheological research is an important tool in reconstructing subsistence, as well as for inferring relevant aspects regarding social behavior in the past. The organization of hunting parties, forms of predation (number and rate of animals slaughtered), and the technology used (tactics and tools) must be taken into account in the identification and classification of hunting methods in prehistory. The archeological recognition of communal hunting reflects an interest in evolutionary terms and their inherent implications for anticipatory capacities, social complexity, and the development of cognitive tools, such as articulated language. Late and Middle Paleolithic faunal assemblages in Europe have produced convincing evidence of communal hunting of large ungulates allowing for the formation of hypotheses concerning the skills of Neanderthals anatomically modern humans as social predators. However, the emergence of this cooperative behavior is not currently understood. Here, faunal analysis, based on traditional/long-established zooarcheological methods, of nearly 25,000 faunal remains from the "bison bone bed" layer of the TD10.2 sub-unit at Gran Dolina, Atapuerca (Spain) is presented. In addition, other datasets related to the archeo-stratigraphy, paleo-landscape, paleo-environmental proxies, lithic assemblage, and ethno-archeological information of communal hunting have been considered in order to adopt a holistic approach to an investigation of the subsistence strategies developed during deposition of the archeological remains. The results indicate a monospecific assemblage heavily dominated by axial bison elements. The abundance of anthropogenic modifications and the anatomical profile are in concordance with early primary access to carcasses and the development of systematic butchering focused on the exploitation of meat and fat for transportation of high-yield elements to somewhere out of the cave. Together with a catastrophic and seasonal mortality pattern, the results indicate the procurement of bison by communal hunting as early as circa 400 kyr. This suggests that the cognitive, social, and technological capabilities required for successful communal hunting were at least fully developed among the pre-Neanderthal paleodeme of Atapuerca during the Lower Paleolithic. Similarly, the early existence of mass communal hunting as a predation technique informs our understanding of the early emergence of predatory skills similar to those exhibited by modern communal hunters.

La zooarqueología es una importante herramienta para la reconstrucción de la subsistencia y también para inferir aspectos relevantes del comportamiento social en el pasado. En este trabajo presentamos el análisis faunístico del llamado "lecho de huesos de bisonte", contenido en la subunidad TD10.2 del yacimiento Gran Dolina (Atapuerca, España). La composición taxonómica y perfil anatómico indican un conjunto monoespecífico fuertemente dominado por elementos del esqueleto axial de bisonte (Bison sp.). Las características y abundancia de las modificaciones antrópicas revelan un acceso primario e inmediato a las carcasas, así como el desarrollo de un procesado carnicero sistemático dirigido a la explotación de la carne y grasa, y a la preparación para el transporte de elementos de alto rendimiento hacia algún lugar fuera de la cavidad. Esas características unidas a un perfil de mortalidad catastrófico y estacional, sugieren la obtención de los bisontes mediante caza comunal. La frecuencia, localización e intensidad de las mordeduras de carnívoro en los restos indica un fuerte saqueo de las carcasas de bisonte una vez abandonadas éstas por los homininos en el yacimiento. La suma de decisiones antrópicas sobre el transporte y el posterior saqueo por carnívoros de los despojos abandonados resulta en un conjunto interpretado como lugar de matanza y procesamiento carnicero de bisontes carroñeados posteriormente por las fieras. Las analogías etnográficas, etnohistóricas y arqueológicas nos han permitido interpretar el "lecho de huesos de bisonte" como cazadero utilizado durante varios eventos estacionales de caza comunal en los que rebaños completos de bisontes fueron sacrificados para ser explotados intensamente por los homininos que ocuparon la cueva. El repetido uso estacional de un punto en el territorio para el desarrollo de tareas específicas muestra ciertas similitudes con el patrón logístico de gestión de los recursos. En el mismo sentido, la existencia temprana de la caza comunal como táctica depredatoria nos informa sobre la emergencia de habilidades cognitivas, tecnológicas y sociales similares a aquellas exhibidas por otros cazadores comunales modernos en un momento tan temprano como el Pleistoceno medio.

Understanding the emergence of modern humans and the disappearance of Neanderthals: Insights from Kaldar Cave (Khorramabad Valley, Western Iran).

Kaldar Cave is a key archaeological site that provides evidence of the Middle to Upper Palaeolithic transition in Iran. Excavations at the site in 2014-2015 led to the discovery of cultural remains generally associated with anatomically modern humans (AMHs) and evidence of a probable Neanderthal-made industry in the basal layers. Attempts have been made to establish a chronology for the site. These include four thermoluminescence (TL) dates for Layer 4, ranging from 23,100 ± 3300 to 29,400 ± 2300 BP, and three AMS radiocarbon dates from charcoal samples belonging to the lower part of the same layer, yielding ages of 38,650-36,750 cal BP, 44,200-42,350 cal BP, and 54,400-46,050 cal BP (all at the 95.4% confidence level). Kaldar Cave is the first well-stratified Late Palaeolithic locality to be excavated in the Zagros which is one of the earliest sites with cultural materials attributed to early AMHs in western Asia. It also offers an opportunity to study the technological differences between the Mousterian and the first Upper Palaeolithic lithic technologies as well as the human behaviour in the region. In this study, we present a detailed description of the newly excavated stratigraphy, quantified results from the lithic assemblages, preliminary faunal remains analyses, geochronologic data, taphonomic aspects, and an interpretation of the regional paleoenvironment.

The diet of the first Europeans from Atapuerca.

Hominin dietary specialization is crucial to understanding the evolutionary changes of craniofacial biomechanics and the interaction of food processing methods' effects on teeth. However, the diet-related dental wear processes of the earliest European hominins remain unknown because most of the academic attention has focused on Neandertals. Non-occlusal dental microwear provides direct evidence of the effect of chewed food particles on tooth enamel surfaces and reflects dietary signals over time. Here, we report for the first time the direct effect of dietary abrasiveness as evidenced by the buccal microwear patterns on the teeth of the Sima del Elefante-TE9 and Gran Dolina-TD6 Atapuerca hominins (1.2-0.8 million years ago - Myr) as compared with other Lower and Middle Pleistocene populations. A unique buccal microwear pattern that is found in Homo antecessor (0.96-0.8 Myr), a well-known cannibal species, indicates dietary practices that are consistent with the consumption of hard and brittle foods. Our findings confirm that the oldest European inhabitants ingested more mechanically-demanding diets than later populations because they were confronted with harsh, fluctuating environmental conditions. Furthermore, the influence of grit-laden food suggests that a high-quality meat diet from butchering processes could have fueled evolutionary changes in brain size.

Diet and environment 1.2 million years ago revealed through analysis of dental calculus from Europe's oldest hominin at Sima del Elefante, Spain.

Sima del Elefante, Atapuerca, Spain contains one of the earliest hominin fragments yet known in Europe, dating to 1.2 Ma. Dental calculus from a hominin molar was removed, degraded and analysed to recover entrapped remains. Evidence for plant use at this time is very limited and this study has revealed the earliest direct evidence for foods consumed in the genus Homo. This comprises starchy carbohydrates from two plants, including a species of grass from the Triticeae or Bromideae tribe, meat and plant fibres. All food was eaten raw, and there is no evidence for processing of the starch granules which are intact and undamaged. Additional biographical detail includes fragments of non-edible wood found adjacent to an interproximal groove suggesting oral hygiene activities, while plant fibres may be linked to raw material processing. Environmental evidence comprises spores, insect fragments and conifer pollen grains which are consistent with a forested environment.

Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins.

A unique assemblage of 28 hominin individuals, found in Sima de los Huesos in the Sierra de Atapuerca in Spain, has recently been dated to approximately 430,000 years ago. An interesting question is how these Middle Pleistocene hominins were related to those who lived in the Late Pleistocene epoch, in particular to Neanderthals in western Eurasia and to Denisovans, a sister group of Neanderthals so far known only from southern Siberia. While the Sima de los Huesos hominins share some derived morphological features with Neanderthals, the mitochondrial genome retrieved from one individual from Sima de los Huesos is more closely related to the mitochondrial DNA of Denisovans than to that of Neanderthals. However, since the mitochondrial DNA does not reveal the full picture of relationships among populations, we have investigated DNA preservation in several individuals found at Sima de los Huesos. Here we recover nuclear DNA sequences from two specimens, which show that the Sima de los Huesos hominins were related to Neanderthals rather than to Denisovans, indicating that the population divergence between Neanderthals and Denisovans predates 430,000 years ago. A mitochondrial DNA recovered from one of the specimens shares the previously described relationship to Denisovan mitochondrial DNAs, suggesting, among other possibilities, that the mitochondrial DNA gene pool of Neanderthals turned over later in their history.

Ontogeny of the maxilla in Neanderthals and their ancestors.

Neanderthals had large and projecting (prognathic) faces similar to those of their putative ancestors from Sima de los Huesos (SH) and different from the retracted modern human face. When such differences arose during development and the morphogenetic modifications involved are unknown. We show that maxillary growth remodelling (bone formation and resorption) of the Devil's Tower (Gibraltar 2) and La Quina 18 Neanderthals and four SH hominins, all sub-adults, show extensive bone deposition, whereas in modern humans extensive osteoclastic bone resorption is found in the same regions. This morphogenetic difference is evident by ∼5 years of age. Modern human faces are distinct from those of the Neanderthal and SH fossils in part because their postnatal growth processes differ markedly. The growth remodelling identified in these fossil hominins is shared with Australopithecus and early Homo but not with modern humans suggesting that the modern human face is developmentally derived.

Genome-wide patterns of selection in 230 ancient Eurasians.

Ancient DNA makes it possible to observe natural selection directly by analysing samples from populations before, during and after adaptation events. Here we report a genome-wide scan for selection using ancient DNA, capitalizing on the largest ancient DNA data set yet assembled: 230 West Eurasians who lived between 6500 and 300 bc, including 163 with newly reported data. The new samples include, to our knowledge, the first genome-wide ancient DNA from Anatolian Neolithic farmers, whose genetic material we obtained by extracting from petrous bones, and who we show were members of the population that was the source of Europe's first farmers. We also report a transect of the steppe region in Samara between 5600 and 300 bc, which allows us to identify admixture into the steppe from at least two external sources. We detect selection at loci associated with diet, pigmentation and immunity, and two independent episodes of selection on height.