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 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.

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.

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.

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.

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.

A mitochondrial genome sequence of a hominin from Sima de los Huesos.

Excavations of a complex of caves in the Sierra de Atapuerca in northern Spain have unearthed hominin fossils that range in age from the early Pleistocene to the Holocene. One of these sites, the 'Sima de los Huesos' ('pit of bones'), has yielded the world's largest assemblage of Middle Pleistocene hominin fossils, consisting of at least 28 individuals dated to over 300,000 years ago. The skeletal remains share a number of morphological features with fossils classified as Homo heidelbergensis and also display distinct Neanderthal-derived traits. Here we determine an almost complete mitochondrial genome sequence of a hominin from Sima de los Huesos and show that it is closely related to the lineage leading to mitochondrial genomes of Denisovans, an eastern Eurasian sister group to Neanderthals. Our results pave the way for DNA research on hominins from the Middle Pleistocene.

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.

Postcranial morphology of the middle Pleistocene humans from Sima de los Huesos, Spain.

Current knowledge of the evolution of the postcranial skeleton in the genus Homo is hampered by a geographically and chronologically scattered fossil record. Here we present a complete characterization of the postcranium of the middle Pleistocene paleodeme from the Sima de los Huesos (SH) and its paleobiological implications. The SH hominins show the following: (i) wide bodies, a plesiomorphic character in the genus Homo inherited from their early hominin ancestors; (ii) statures that can be found in modern human middle-latitude populations that first appeared 1.6-1.5 Mya; and (iii) large femoral heads in some individuals, a trait that first appeared during the middle Pleistocene in Africa and Europe. The intrapopulational size variation in SH shows that the level of dimorphism was similar to modern humans (MH), but the SH hominins were less encephalized than Neandertals. SH shares many postcranial anatomical features with Neandertals. Although most of these features appear to be either plesiomorphic retentions or are of uncertain phylogenetic polarity, a few represent Neandertal apomorphies. Nevertheless, the full suite of Neandertal-derived features is not yet present in the SH population. The postcranial evidence is consistent with the hypothesis based on the cranial morphology that the SH hominins are a sister group to the later Neandertals. Comparison of the SH postcranial skeleton to other hominins suggests that the evolution of the postcranium occurred in a mosaic mode, both at a general and at a detailed level.

Ancient genomes link early farmers from Atapuerca in Spain to modern-day Basques.

The consequences of the Neolithic transition in Europe--one of the most important cultural changes in human prehistory--is a subject of great interest. However, its effect on prehistoric and modern-day people in Iberia, the westernmost frontier of the European continent, remains unresolved. We present, to our knowledge, the first genome-wide sequence data from eight human remains, dated to between 5,500 and 3,500 years before present, excavated in the El Portalón cave at Sierra de Atapuerca, Spain. We show that these individuals emerged from the same ancestral gene pool as early farmers in other parts of Europe, suggesting that migration was the dominant mode of transferring farming practices throughout western Eurasia. In contrast to central and northern early European farmers, the Chalcolithic El Portalón individuals additionally mixed with local southwestern hunter-gatherers. The proportion of hunter-gatherer-related admixture into early farmers also increased over the course of two millennia. The Chalcolithic El Portalón individuals showed greatest genetic affinity to modern-day Basques, who have long been considered linguistic and genetic isolates linked to the Mesolithic whereas all other European early farmers show greater genetic similarity to modern-day Sardinians. These genetic links suggest that Basques and their language may be linked with the spread of agriculture during the Neolithic. Furthermore, all modern-day Iberian groups except the Basques display distinct admixture with Caucasus/Central Asian and North African groups, possibly related to historical migration events. The El Portalón genomes uncover important pieces of the demographic history of Iberia and Europe and reveal how prehistoric groups relate to modern-day people.

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.

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.

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.

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.

No known hominin species matches the expected dental morphology of the last common ancestor of Neanderthals and modern humans.

A central problem in paleoanthropology is the identity of the last common ancestor of Neanderthals and modern humans ([N-MH]LCA). Recently developed analytical techniques now allow this problem to be addressed using a probabilistic morphological framework. This study provides a quantitative reconstruction of the expected dental morphology of the [N-MH]LCA and an assessment of whether known fossil species are compatible with this ancestral position. We show that no known fossil species is a suitable candidate for being the [N-MH]LCA and that all late Early and Middle Pleistocene taxa from Europe have Neanderthal dental affinities, pointing to the existence of a European clade originated around 1 Ma. These results are incongruent with younger molecular divergence estimates and suggest at least one of the following must be true: (i) European fossils and the [N-MH]LCA selectively retained primitive dental traits; (ii) molecular estimates of the divergence between Neanderthals and modern humans are underestimated; or (iii) phenotypic divergence and speciation between both species were decoupled such that phenotypic differentiation, at least in dental morphology, predated speciation.

Early Pleistocene human hand phalanx from the Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain).

In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante cave site (TE - Sierra de Atapuerca, Spain), ascribed to Homo sp., is presented and comparatively described in the context of the evolution of the genus Homo. The ATE9-2 specimen is especially important because of the paucity of hand bones in the human fossil record during the Early Pleistocene. The morphological and metrical analyses of the phalanx ATE9-2 indicate that there are no essential differences between it and comparator fossil specimens for the genus Homo after 1.3 Ma (millions of years ago). Similar to Sima de los Huesos and Neandertal specimens, ATE9-2 is a robust proximal hand phalanx, probably reflecting greater overall body robusticity in these populations or a higher gracility in modern humans. The age of level TE9 from Sima del Elefante and morphological and metrical studies of ATE9-2 suggest that the morphology of the proximal hand phalanges and, thus, the morphology of the hand could have remained stable over the last 1.2-1.3 Ma. Taking into account the evidence recently provided by a metacarpal from Kaitio (Kenya) from around 1.42 Ma, we argue that modern hand morphology is present in the genus Homo subsequent to Homo habilis.

The lithic industry of Sima del Elefante (Atapuerca, Burgos, Spain) in the context of Early and Middle Pleistocene technology in Europe.

This paper presents the lithic assemblages documented at Sima del Elefante (TE) and their importance in the context of the Early and Middle Pleistocene human occupation of Europe. We also study changes in human behaviour within the context of the palaeoenvironmental evolution of the Sierra de Atapuerca. This site has characteristics that are of great value for the study of human evolution. The lower levels of TE (Units TE7-TE14) are an essential reference for understanding the early stages of the colonization of Europe. The TE9c level has provided stone tools (Mode 1), faunal remains, and human fossils dated to 1.22 Ma (millions of years ago). Moreover, this is one of the few European sites with a stratigraphic sequence that includes remains of human occupations predating the Jaramillo subchron (Early Pleistocene) and from the Late Middle Pleistocene (Units TE18-TE19). Despite this, the presence of archaeologically sterile units (TE15-17) prevents us from establishing a continuous relationship between the Early and Middle Pleistocene human settlements and, consequently, between their technological and behavioural differences. We can, however compare the technological and palaeoeconomic strategies adopted by different species of hominins during two key phases of the occupation of Europe.

The medial pterygoid tubercle in the Atapuerca Early and Middle Pleistocene mandibles: evolutionary implications.

Numerous studies have attempted to identify the presence of uniquely derived (autoapomorphic) Neandertal features. Here, we deal with the medial pterygoid tubercle (MTP), which is usually present on the internal face of the ascending ramus of Neandertal specimens. Our study stems from the identification of a hypertrophied tubercle in ATD6-96, an Early Pleistocene mandible recovered from the TD6 level of the Atapuerca-Gran Dolina site and attributed to Homo antecessor. Our review of the literature and study of numerous original fossil specimens and high quality replicas confirm that the MTP occurs at a high frequency in Neandertals (ca. 89%) and is also present in over half (ca. 55%) of the Middle Pleistocene Sima de los Huesos (SH) hominins. In contrast, it is generally absent or minimally developed in other extinct hominins, but can be found in variable frequencies (

Luminescence dating and palaeomagnetic age constraint on hominins from Sima de los Huesos, Atapuerca, Spain.

Establishing a reliable chronology on the extensive hominin remains at Sima de los Huesos is critical for an improved understanding of the complex evolutionary histories and phylogenetic relationships of the European Middle Pleistocene hominin record. In this study, we use a combination of 'extended-range' luminescence dating techniques and palaeomagnetism to provide new age constraint on sedimentary infills that are unambiguously associated with the Sima fossil assemblage. Post-infrared-infrared stimulated luminescence (pIR-IR) dating of K-feldspars and thermally transferred optically stimulated luminescence (TT-OSL) dating of individual quartz grains provide weighted mean ages of 433 ± 15 ka (thousands of years) and 416 ± 19 ka, respectively, for allochthonous sedimentary horizons overlying the hominin-bearing clay breccia. The six replicate luminescence ages obtained for this deposit are reproducible and provide a combined minimum age estimate of 427 ± 12 ka for the underlying hominin fossils. Palaeomagnetic directions for the luminescence dated sediment horizon and underlying fossiliferous clays display exclusively normal polarities. These findings are consistent with the luminescence dating results and confirm that the hominin fossil horizon accumulated during the Brunhes Chron, i.e., within the last 780 ka. The new bracketing age constraint for the Sima hominins is in broad agreement with radiometrically dated Homo heidelbergensis fossil sites, such as Mauer and Arago, and suggests that the split of the H. neanderthalensis and H. sapiens lineages took place during the early Middle Pleistocene. More widespread numerical dating of key Early and Middle Pleistocene fossil sites across Europe is needed to test and refine competing models of hominin evolution. The new luminescence chronologies presented in this study demonstrate the versatility of TT-OSL and pIR-IR techniques and the potential role they could play in helping to refine evolutionary histories over Middle Pleistocene timescales.

The first hominin of Europe.

The earliest hominin occupation of Europe is one of the most debated topics in palaeoanthropology. However, the purportedly oldest of the Early Pleistocene sites in Eurasia lack precise age control and contain stone tools rather than human fossil remains. Here we report the discovery of a human mandible associated with an assemblage of Mode 1 lithic tools and faunal remains bearing traces of hominin processing, in stratigraphic level TE9 at the site of the Sima del Elefante, Atapuerca, Spain. Level TE9 has been dated to the Early Pleistocene (approximately 1.2-1.1 Myr), based on a combination of palaeomagnetism, cosmogenic nuclides and biostratigraphy. The Sima del Elefante site thus emerges as the oldest, most accurately dated record of human occupation in Europe, to our knowledge. The study of the human mandible suggests that the first settlement of Western Europe could be related to an early demographic expansion out of Africa. The new evidence, with previous findings in other Atapuerca sites (level TD6 from Gran Dolina), also suggests that a speciation event occurred in this extreme area of the Eurasian continent during the Early Pleistocene, initiating the hominin lineage represented by the TE9 and TD6 hominins.