title: Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins.
abstract: 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.
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Matthias
authors.initials: M
authors.lastname: Meyer
authors.affiliation: Centro de Investigación Sobre la Evolución y Comportamiento Humanos, Universidad Complutense de Madrid-Instituto de Salud Carlos III, 28029 Madrid, Spain.|Departamento de Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain.
authors.forename: Juan-Luis
authors.initials: JL
authors.lastname: Arsuaga
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Cesare
authors.initials: C
authors.lastname: de Filippo
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Sarah
authors.initials: S
authors.lastname: Nagel
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Ayinuer
authors.initials: A
authors.lastname: Aximu-Petri
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Birgit
authors.initials: B
authors.lastname: Nickel
authors.affiliation: Centro de Investigación Sobre la Evolución y Comportamiento Humanos, Universidad Complutense de Madrid-Instituto de Salud Carlos III, 28029 Madrid, Spain.|Área de Paleontología, Departamento de Geografía y Geología, Universidad de Alcalá, Alcalá de Henares, 28871 Madrid, Spain.
authors.forename: Ignacio
authors.initials: I
authors.lastname: Martínez
authors.affiliation: Centro de Investigación Sobre la Evolución y Comportamiento Humanos, Universidad Complutense de Madrid-Instituto de Salud Carlos III, 28029 Madrid, Spain.|Área de Paleontología, Departamento de Geografía y Geología, Universidad de Alcalá, Alcalá de Henares, 28871 Madrid, Spain.
authors.forename: Ana
authors.initials: A
authors.lastname: Gracia
authors.affiliation: Centro Nacional de Investigación sobre la Evolución Humana, Paseo Sierra de Atapuerca, 09002 Burgos, Spain.
authors.forename: José María
authors.initials: JM
authors.lastname: Bermúdez de Castro
authors.affiliation: Institut Català de Paleoecologia Humana i Evolució Social, C/Marcel·lí Domingo s/n (Edifici W3), Campus Sescelades, 43007 Tarragona, Spain.|Àrea de Prehistòria, Departament d'Història i Història de l'Art, Universitat Rovira i Virgili, Facultat de Lletres, Avinguda de Catalunya, 35, 43002 Tarragona, Spain.
authors.forename: Eudald
authors.initials: E
authors.lastname: Carbonell
authors.affiliation: Department of Anthropology, University of Toronto, 19 Russell Street, Toronto, Ontario M5S 2S2, Canada.
authors.forename: Bence
authors.initials: B
authors.lastname: Viola
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Janet
authors.initials: J
authors.lastname: Kelso
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Kay
authors.initials: K
authors.lastname: Prüfer
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Svante
authors.initials: S
authors.lastname: Pääbo
mesh_terms: D000483:Alleles; D000818:Animals; D004272:DNA, Mitochondrial / Q000235:genetics; D005580:Fossils; D054629:Genome, Mitochondrial / Q000235:genetics; D015186:Hominidae / Q000145:classification / Q000235:genetics*; D008297:Male; D059125:Neanderthals / Q000145:classification / Q000235:genetics; D010802:Phylogeny; D016415:Sequence Alignment; D013030:Spain
chemical_list: D004272:DNA, Mitochondrial