title: A revised timescale for human evolution based on ancient mitochondrial genomes.
abstract: BACKGROUND
  Recent analyses of de novo DNA mutations in modern humans have suggested a nuclear substitution rate that is approximately half that of previous estimates based on fossil calibration. This result has led to suggestions that major events in human evolution occurred far earlier than previously thought.
  
  
  RESULTS
  Here, we use mitochondrial genome sequences from ten securely dated ancient modern humans spanning 40,000 years as calibration points for the mitochondrial clock, thus yielding a direct estimate of the mitochondrial substitution rate. Our clock yields mitochondrial divergence times that are in agreement with earlier estimates based on calibration points derived from either fossils or archaeological material. In particular, our results imply a separation of non-Africans from the most closely related sub-Saharan African mitochondrial DNAs (haplogroup L3) that occurred less than 62-95 kya.
  
  
  CONCLUSIONS
  Though single loci like mitochondrial DNA (mtDNA) can only provide biased estimates of population divergence times, they can provide valid upper bounds. Our results exclude most of the older dates for African and non-African population divergences recently suggested by de novo mutation rate estimates in the nuclear genome.
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103 Germany.|CAS-MPS Joint Laboratory for Human Evolution and Archaeometry, Institute of Vertebrate Paleontology and Paleoanthropology of Chinese Academy of Sciences, Beijing 100044, P.R. China.
authors.forename: Qiaomei
authors.initials: Q
authors.lastname: Fu
authors.affiliation: Institute for Archaeological Sciences, University of Tübingen, Rümelinstr. 23, Tübingen, Germany.
authors.forename: Alissa
authors.initials: A
authors.lastname: Mittnik
authors.affiliation: Department of Biology, Emory University, Atlanta, GA, 30322 USA.
authors.forename: Philip L F
authors.initials: PLF
authors.lastname: Johnson
authors.affiliation: Institute for Archaeological Sciences, University of Tübingen, Rümelinstr. 23, Tübingen, Germany.|Laboratoire de Paléoanthropologie, École Pratique des Hautes Études, UMR 5199 PACEA, CNRS-Université de Bordeaux, Bordeaux, France.
authors.forename: Kirsten
authors.initials: K
authors.lastname: Bos
authors.affiliation: Dipartimento di Biologia Evoluzionistica, Università di Firenze, Firenze, Italy.
authors.forename: Martina
authors.initials: M
authors.lastname: Lari
authors.affiliation: Palaeogenetics Group, Institute for Anthropology, Johannes Gutenberg-University, Saarstrasse 21, D-55099 Mainz, Germany.
authors.forename: Ruth
authors.initials: R
authors.lastname: Bollongino
authors.affiliation: ShandongMuseum, 11899 Jing 10th Road, Jinan, Shandong 250014, P. R. China.
authors.forename: Chengkai
authors.initials: C
authors.lastname: Sun
authors.affiliation: LVR-Landesmuseum Bonn, Bachstrasse 5-9, D-53115 Bonn, Germany.|Department of Prehistoric and Protohistoric Archaeology, Institute for Archaeology and Cultural Anthropology, University of Bonn, Regina-Pacis-Weg 7, 53113 Bonn, Germany.
authors.forename: Liane
authors.initials: L
authors.lastname: Giemsch
authors.affiliation: LVR-Landesmuseum Bonn, Bachstrasse 5-9, D-53115 Bonn, Germany.
authors.forename: Ralf
authors.initials: R
authors.lastname: Schmitz
authors.affiliation: Palaeogenetics Group, Institute for Anthropology, Johannes Gutenberg-University, Saarstrasse 21, D-55099 Mainz, Germany.
authors.forename: Joachim
authors.initials: J
authors.lastname: Burger
authors.affiliation: Università degli Studi di Siena Dip. di Scienze Fisiche, della Terra e dell'Ambiente U.R. Ecologia Preistorica Via Laterina, 8 - 53100 Siena, Italy.
authors.forename: Anna Maria
authors.initials: AM
authors.lastname: Ronchitelli
authors.affiliation: Università di Firenze Dipartimento di Scienze dell'Antichità, Medioevo e Rinascimento e Linguistica, Piazza Brunelleschi, 3-4 - 50121 Firenze, Italy.
authors.forename: Fabio
authors.initials: F
authors.lastname: Martini
authors.affiliation: Dipartimento di Scienze Archeologiche Università di Pisa via Galvani 1, 56126 Pisa.
authors.forename: Renata G
authors.initials: RG
authors.lastname: Cremonesi
authors.affiliation: Department of Anthropology, Faculty of Science, Masaryk University, Vinařská 5, 603 00 Brno.|Institute of Archaeology, Academy of Science of the Czech Republic, Kralovopolska 147, 612 00 Brno, Czech Republic.
authors.forename: Jiří
authors.initials: J
authors.lastname: Svoboda
authors.affiliation: Human Genetics Department, Medical Faculty, University of Tübingen, 72070 Tübingen, Germany.
authors.forename: Peter
authors.initials: P
authors.lastname: Bauer
authors.affiliation: Dipartimento di Biologia Evoluzionistica, Università di Firenze, Firenze, Italy.
authors.forename: David
authors.initials: D
authors.lastname: Caramelli
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103 Germany.
authors.forename: Sergi
authors.initials: S
authors.lastname: Castellano
authors.affiliation: Broad Institute of MIT and Harvard, Cambridge, MA, 02142 USA.|Department of Genetics, Harvard Medical School, Boston, MA, 02115 USA.
authors.forename: David
authors.initials: D
authors.lastname: Reich
authors.affiliation: Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103 Germany.
authors.forename: Svante
authors.initials: S
authors.lastname: Pääbo
authors.affiliation: Institute for Archaeological Sciences, University of Tübingen, Rümelinstr. 23, Tübingen, Germany.
authors.forename: Johannes
authors.initials: J
authors.lastname: Krause
mesh_terms: D001483:Base Sequence; D001499:Bayes Theorem; D003710:Demography; D019143:Evolution, Molecular; D005580:Fossils; D015894:Genome, Human / Q000235:genetics*; D054629:Genome, Mitochondrial / Q000235:genetics*; D006239:Haplotypes / Q000235:genetics; D059014:High-Throughput Nucleotide Sequencing; D006801:Humans; D016014:Linear Models; D008957:Models, Genetic; D008969:Molecular Sequence Data; D010802:Phylogeny; D013997:Time Factors