Who We Are and How We Got Here

by David Reich

  24. Ibid.

  25. Ibid; M. Meyer et al., “A High-Coverage Genome Sequence from an Archaic Denisovan Individual,” Science 338 (2012): 222–26; J. D. Wall et al., “Higher Levels of Neanderthal Ancestry in East Asians Than in Europeans,” Genetics 194 (2013): 199–209.

  26. Q. Fu et al., “The Genetic History of Ice Age Europe,” Nature 534 (2016): 200–5.

  27. I. Lazaridis et al., “Genomic Insights into the Origin of Farming in the Ancient Near East,” Nature 536 (2016): 419–24.

  28. Trinkaus et al., “An Early Modern Human.”

  29. Q. Fu et al., “An Early Modern Human from Romania with a Recent Neanderthal Ancestor,” Nature 524 (2015): 216–19.

  30. N. Teyssandier, F. Bon, and J.-G. Bordes, “Within Projectile Range: Some Thoughts on the Appearance of the Aurignacian in Europe,” Journal of Anthropological Research 66 (2010): 209–29; P. Mellars, “Archeology and the Dispersal of Modern Humans in Europe: Deconstructing the ‘Aurignacian,’ ” Evolutionary Anthropology 15 (2006): 167–82.

  31. M. Currat and L. Excoffier, “Strong Reproductive Isolation Between Humans and Neanderthals Inferred from Observed Patterns of Introgression,” Proceedings of the National Academy of Sciences of the U.S.A. 108 (2011): 15129–34.

  32. S. Sankararaman et al., “The Genomic Landscape of Neanderthal Ancestry in Present-Day Humans,” Nature 507 (2014): 354–57; B. Vernot and J. M. Akey, “Resurrecting Surviving Neandertal Lineages from Modern Human Genomes,” Science 343 (2014): 1017-21.

  33. N. Patterson et al., “Genetic Evidence for Complex Speciation of Humans and Chimpanzees,” Nature 441 (2006): 1103–8.

  34. Ibid; R. Burgess and Z. Yang, “Estimation of Hominoid Ancestral Population Sizes Under Bayesian Coalescent Models Incorporating Mutation Rate Variation and Sequencing Errors,” Molecular Biology and Evolution 25 (2008): 1975–94.

  35. J. A. Coyne and H. A. Orr, “Two Rules of Speciation,” in Speciation and Its Consequences, ed. Daniel Otte and John A. Endler (Sunderland, MA: Sinauer Associates, 1989), 180–207.

  36. P. K. Tucker et al., “Abrupt Cline for Sex-Chromosomes in a Hybrid Zone Between Two Species of Mice,” Evolution 46 (1992): 1146–63.

  37. H. Li and R. Durbin, “Inference of Human Population History from Individual Whole-Genome Sequences,” Nature 475 (2011): 493–96.

  38. T. Mailund et al., “A New Isolation with Migration Model Along Complete Genomes Infers Very Different Divergence Processes Among Closely Related Great Ape Species,” PLoS Genetics 8 (2012): e1003125.

  39. J. Y. Dutheil et al., “Strong Selective Sweeps on the X Chromosome in the Human-Chimpanzee Ancestor Explain Its Low Divergence,” PLoS Genetics 11 (2015): e1005451.

  40. Sankararaman et al., “Genomic Landscape”; B. Jégou et al., “Meiotic Genes Are Enriched in Regions of Reduced Archaic Ancestry,” Molecular Biology and Evolution 34 (2017): 1974–80.

  41. Q. Fu et al., “Ice Age Europe.”

  42. I. Juric, S. Aeschbacher, and G. Coop, “The Strength of Selection Against Neanderthal Introgression,” PLoS Genetics 12 (2016): e1006340; K. Harris and R. Nielsen, “The Genetic Cost of Neanderthal Introgression,” Genetics 203 (2016): 881–91.

  43. G. Bhatia et al., “Genome-Wide Scan of 29,141 African Americans Finds No Evidence of Directional Selection Since Admixture,” American Journal of Human Genetics 95 (2014): 437–44.

  44. Johann G. Fichte, Grundlage der gesamten Wissenschaftslehre (Jena, Germany: Gabler, 1794).

  3 Ancient DNA Opens the Floodgates

  1. J. Krause et al., “Neanderthals in Central Asia and Siberia,” Nature 449 (2007): 902–4.

  2. J. Krause et al., “The Complete Mitochondrial DNA Genome of an Unknown Hominin from Southern Siberia,” Nature 464 (2010): 894-97.

  3. C. Posth et al., “Deeply Divergent Archaic Mitochondrial Genome Provides Lower Time Boundary for African Gene Flow into Neanderthals,” Nature Communications 8 (2017): 16046.

  4. Krause et al., “Unknown Hominin.”

  5. D. Reich et al., “Genetic History of an Archaic Hominin Group from Denisova Cave in Siberia,” Nature 468 (2010): 1053–60.

  6. K. Prüfer et al., “The Complete Genome Sequence of a Neanderthal from the Altai Mountains,” Nature (2013): doi: 10.1038/nature 12886.

  7. Jerry A. Coyne and H. Allen Orr, Speciation (Sunderland, MA: Sinauer Associates, 2004).

  8. S. Sankararaman, S. Mallick, N. Patterson, and D. Reich, “The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans,” Current Biology 26 (2016): 1241–47.

  9. P. Moorjani et al., “A Genetic Method for Dating Ancient Genomes Provides a Direct Estimate of Human Generation Interval in the Last 45,000 Years,” Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 5652–7.

  10. Sankararaman et al., “Combined Landscape.”

  11. D. Reich et al., “Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania,” American Journal of Human Genetics 89 (2011): 516–28.

  12. Q. Fu et al., “DNA Analysis of an Early Modern Human from Tianyuan Cave, China,” Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 2223–27; M. Yang et al., “40,000-Year-Old Individual from Asia Provides Insight into Early Population Structure in Eurasia,” Current Biology 27 (2017): 3202–8.

  13. Prüfer et al., “Complete Genome.”

  14. C. B. Stringer and I. Barnes, “Deciphering the Denisovans,” Proceedings of the National Academy of Sciences of the U.S.A. 112 (2015): 15542–43.

  15. G. A. Wagner et al., “Radiometric Dating of the Type-Site for Homo Heidelbergensis at Mauer, Germany,” Proceedings of the National Academy of Sciences of the U.S.A. 107 (2010): 19726–30.

  16. C. Stringer, “The Status of Homo heidelbergensis (Schoetensack 1908),” Evolutionary Anthropology 21 (2012): 101–7.

  17. A. Brumm et al., “Age and Context of the Oldest Known Hominin Fossils from Flores,” Nature 534 (2016): 249–53.

  18. Reich et al., “Denisova Admixture.”

  19. Prüfer et al., “Complete Genome.”

  20. Ibid.; Sankararaman et al., “Combined Landscape.”

  21. E. Huerta-Sánchez et al., “Altitude Adaptation in Tibetans Caused by Introgression of Denisovan-like DNA,” Nature 512 (2014): 194–97.

  22. F. H. Chen et al., “Agriculture Facilitated Permanent Human Occupation of the Tibetan Plateau After 3600 B.P.,” Science 347 (2015): 248–50.

  23. S. Sankararaman et al., “The Genomic Landscape of Neanderthal Ancestry in Present-Day Humans,” Nature 507 (2014): 354–57; B. Vernot and J. M. Akey, “Resurrecting Surviving Neandertal Lineages from Modern Human Genomes,” Science 343 (2014): 1017–21.

  24. Prüfer et al., “Complete Genome.”

  25. G. P. Rightmire, “Homo erectus and Middle Pleistocene Hominins: Brain Size, Skull Form, and Species Recognition,” Journal of Human Evolution 65 (2013): 223–52.

  26. M. Martinón-Torres et al., “Dental Evidence on the Hominin Dispersals During the Pleistocene,” Proceedings of the National Academy of Sciences of the U.S.A. 104 (2007): 13279–82; M. Martinón-Torres, R. Dennell, and J. M. B. de Castro, “The Denisova Hominin Need Not Be an Out of Africa Story,” Journal of Human Evolution 60 (2011): 251–55; J. M. B. de Castro and M. Martinón-Torres, “A New Model for the Evolution of the Human Pleistocene Populations of Europe,” Quaternary International 295 (2013): 102–12.

  27. De Castro and Martinón-Torres, “A New Model.”

  28. J. L. Arsuaga et al., “Neandertal Roots: Cranial and Chronological Evidence from Sima de los Huesos,” Science 344 (2014): 1358–63; M. Meyer et al., “A Mitochondrial Genome Sequence of a Hominin from Sima de los Huesos,” Nature 505 (2014): 403–6.

  29. M. Meyer et al., “Nuclear DNA Sequences from the Middle Pleistocene Sima de los Huesos Hominins,” Nature 531 (2016): 504–7.

  30. Meyer et al., “A Mitochondrial Genome”; Meyer et al., “Nuclear DNA Sequences.”

  31. Krause
et al., “Unknown Hominin”; Reich et al., “Genetic History.”

  32. Posth et al., “Deeply Divergent Archaic.”

  33. Ibid.

  34. Prüfer et al., “Complete Genome.”

  35. S. McBrearty and A. S. Brooks, “The Revolution That Wasn’t: A New Interpretation of the Origin of Modern Human Behavior,” Journal of Human Evolution 39 (2000): 453–563.

  36. M. Kuhlwilm et al., “Ancient Gene Flow from Early Modern Humans into Eastern Neanderthals,” Nature 530 (2016): 429–33.

  4 Humanity’s Ghosts

  1. Charles R. Darwin, On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (London: John Murray, 1859).

  2. C. Becquet et al., “Genetic Structure of Chimpanzee Populations,” PLoS Genetics 3 (2007): e66.

  3. R. E. Green et al., “A Draft Sequence of the Neandertal Genome,” Science 328 (2010): 710–22.

  4. N. J. Patterson et al., “Ancient Admixture in Human History,” Genetics 192 (2012): 1065–93.

  5. Ernst Mayr, Systematics and the Origin of Species from the Viewpoint of a Zoologist (New York: Columbia University Press, 1942).

  6. J. K. Pickrell and D. Reich, “Toward a New History and Geography of Human Genes Informed by Ancient DNA,” Trends in Genetics 30 (2014): 377–89.

  7. A. R. Templeton, “Biological Races in Humans,” Studies in History and Philosophy of Biological and Biomedical Science 44 (2013): 262–71.

  8. M. Raghavan et al., “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans,” Nature 505 (2014): 87–91.

  9. I. Lazaridis et al., “Ancient Human Genomes Suggest Three Ancestral Populations for Present-Day Europeans,” Nature 513 (2014): 409–13.

  10. I. Lazaridis et al., “Genomic Insights into the Origin of Farming in the Ancient Near East,” Nature 536 (2016): 419–24.

  11. Ibid.

  12. F. Broushaki et al., “Early Neolithic Genomes from the Eastern Fertile Crescent,” Science 353 (2016): 499–503; E. R. Jones et al., “Upper Palaeolithic Genomes Reveal Deep Roots of Modern Eurasians,” Nature Communications 6 (2015): 8912.

  13. B. M. Henn et al., “Genomic Ancestry of North Africans Supports Back-to-Africa Migrations,” PLoS Genetics 8 (2012): e1002397.

  14. Lazaridis et al., “Genomic Insights.”

  15. O. Bar-Yosef, “Pleistocene Connections Between Africa and Southwest Asia: An Archaeological Perspective,” African Archaeological Review 5 (1987): 29–38.

  16. Lazaridis et al., “Genomic Insights.”

  17. Lazaridis et al., “Ancient Human Genomes.”

  18. Q. Fu et al., “The Genetic History of Ice Age Europe,” Nature 534 (2016): 200–5.

  19. Q. Fu et al., “Genome Sequence of a 45,000-Year-Old Modern Human from Western Siberia,” Nature 514 (2014): 445–49.

  20. Q. Fu et al., “An Early Modern Human from Romania with a Recent Neanderthal Ancestor,” Nature 524 (2015): 216–19.

  21. F. G. Fedele, B. Giaccio, and I. Hajdas, “Timescales and Cultural Process at 40,000 BP in the Light of the Campanian Ignimbrite Eruption, Western Eurasia,” Journal of Human Evolution 55 (2008): 834–57; A. Costa et al., “Quantifying Volcanic Ash Dispersal and Impact of the Campanian Ignimbrite Super-Eruption,” Geophysical Research Letters 39 (2012): L10310.

  22. Fedele et al., “Timescales and Cultural Process.”

  23. A. Seguin-Orlando et al., “Genomic Structure in Europeans Dating Back at Least 36,200 Years,” Science 346 (2014): 1113–18.

  24. Fu et al., “Ice Age Europe.”

  25. Andreas Maier, The Central European Magdalenian: Regional Diversity and Internal Variability (Dordrecht, The Netherlands: Springer, 2015).

  26. Fu et al., “Ice Age Europe.”

  27. N. A. Rosenberg et al., “Clines, Clusters, and the Effect of Study Design on the Inference of Human Population Structure,” PLoS Genetics 1 (2005): e70; G. Coop et al., “The Role of Geography in Human Adaptation,” PLoS Genetics 5 (2009): e1000500.

  28. Q. Fu et al., “DNA Analysis of an Early Modern Human from Tianyuan Cave, China,” Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 2223–27.

  29. Fu et al., “Recent Neanderthal Ancestor”; W. Haak et al., “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe,” Nature 522 (2015): 207–11.

  30. R. Pinhasi et al., “Optimal Ancient DNA Yields from the Inner Ear Part of the Human Petrous Bone,” PLoS One 10 (2015): e0129102.

  31. Lazaridis et al., “Genomic Insights.”

  32. Ibid.; Broushaki et al., “Early Neolithic Genomes.”

  33. I. Olalde et al., “Derived Immune and Ancestral Pigmentation Alleles in a 7,000-Year-Old Mesolithic European,” Nature 507 (2014): 225–28.

  34. I. Mathieson et al., “Genome-Wide Patterns of Selection in 230 Ancient Eurasians,” Nature 528 (2015): 499–503.

  35. I. Mathieson et al., “The Genomic History of Southeastern Europe,” bioRxiv (2017): doi.org/​10.1101/​135616.

  36. Haak et al., “Massive Migration”; M. E. Allentoft et al., “Population Genomics of Bronze Age Eurasia,” Nature 522 (2015): 167–72.

  37. Templeton, “Biological Races.”

  5 The Making of Modern Europe

  1. B. Bramanti et al., “Genetic Discontinuity Between Local Hunter-Gatherers and Central Europe’s First Farmers,” Science 326 (2009): 137–40.

  2. A. Keller et al., “New Insights into the Tyrolean Iceman’s Origin and Phenotype as Inferred by Whole-Genome Sequencing,” Nature Communications 3 (2012): 698.

  3. W. Muller et al., “Origin and Migration of the Alpine Iceman,” Science 302 (2003): 862–66.

  4. P. Skoglund et al., “Origins and Genetic Legacy of Neolithic Farmers and Hunter-Gatherers in Europe,” Science 336 (2012): 466–69.

  5. Albert J. Ammerman and Luigi Luca Cavalli-Sforza, The Neolithic Transition and the Genetics of Populations in Europe (Princeton, NJ: Princeton University Press, 1984).

  6. N. J. Patterson et al., “Ancient Admixture in Human History,” Genetics 192 (2012): 1065–93.

  7. M. Raghavan et al., “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans,” Nature (2013): doi: 10.1038/nature12736.

  8. I. Lazaridis et al., “Ancient Human Genomes Suggest Three Ancestral Populations for Present-Day Europeans,” Nature 513 (2014): 409–13.

  9. C. Gamba et al., “Genome Flux and Stasis in a Five Millennium Transect of European Prehistory,” Nature Communications 5 (2014): 5257; M. E. Allentoft et al., “Population Genomics of Bronze Age Eurasia,” Nature 522 (2015): 167–72; W. Haak et al., “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe,” Nature 522 (2015): 207–11; I. Mathieson et al., “Genome-Wide Patterns of Selection in 230 Ancient Eurasians,” Nature 528 (2015): 499–503.

  10. Luigi Luca Cavalli-Sforza, Paolo Menozzi, and Alberto Piazza, The History and Geography of Human Genes (Princeton, NJ: Princeton University Press, 1994).

  11. Haak et al., “Massive Migration”; Mathieson et al., “Genome-Wide Patterns.”

  12. Q. Fu et al., “The Genetic History of Ice Age Europe,” Nature 534 (2016): 200–5.

  13. I. Mathieson, “The Genomic History of Southeastern Europe,” bioRxiv (2017): doi.org/​10.1101/​135616.

  14. K. Douka et al., “Dating Knossos and the Arrival of the Earliest Neolithic in the Southern Aegean,” Antiquity 91 (2017): 304–21.

  15. Haak et al., “Massive Migration”; M. Lipson et al., “Parallel Palaeogenomic Transects Reveal Complex Genetic History of Early European Farmers,” Nature 551 (2017): 368–72.

  16. Colin Renfrew, Before Civilization: The Radiocarbon Revolution and Prehistoric Europe (London: Jonathan Cape, 1973).

  17. Marija Gimbutas, The Prehistory of Eastern Europe, Part I: Mesolithic, Neolithic and Copper Age Cultures in Russia and the Baltic Area (American School of Prehistoric Research, Harvard University, Bulletin No. 20) (Ca
mbridge, MA: Peabody Museum, 1956).

  18. David W. Anthony, The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World (Princeton, NJ: Princeton University Press, 2007).

  19. Ibid.

  20. Ibid.

  21. Haak et al., “Massive Migration.”

  22. Ibid.; I. Lazaridis et al., “Genomic Insights into the Origin of Farming in the Ancient Near East,” Nature 536 (2016): 419–24.

  23. M. Ivanova, “Kaukasus Und Orient: Die Entstehung des ‘Maikop-Phänomens’ im 4. Jahrtausend v. Chr.,” Praehistorische Zeitschrift 87 (2012): 1–28.

  24. Haak et al., “Massive Migration”; Allentoft et al., “Bronze Age Eurasia.”

  25. Ibid.

  26. G. Kossinna, “Die Deutsche Ostmark: Ein Heimatboden der Germanen,” Berlin (1919).

  27. B. Arnold, “The Past as Propaganda: Totalitarian Archaeology in Nazi Germany,” Antiquity 64 (1990): 464–78.

  28. H. Härke, “The Debate on Migration and Identity in Europe,” Antiquity 78 (2004): 453–56.

  29. V. Heyd, “Kossinna’s Smile,” Antiquity 91 (2017): 348–59; M. Vander Linden, “Population History in Third-Millennium-BC Europe: Assessing the Contribution of Genetics,” World Archaeology 48 (2016): 714–28; N. N. Johannsen, G. Larson, D. J. Meltzer, and M. Vander Linden, “A Composite Window into Human History,” Science 356 (2017): 1118–20.

  30. Vere Gordon Childe, The Aryans: A Study of Indo-European Origins (London and New York: K. Paul, Trench, Trubner and Co. and Alfred A. Knopf, 1926).

  31. Härke, “Debate on Migration and Identity.”

  32. Peter Bellwood, First Migrants: Ancient Migration in Global Perspective (Chichester, West Sussex, UK / Malden, MA: Wiley-Blackwell, 2013).

  33. Colin McEvedy and Richard Jones, Atlas of World Population History (Harmondsworth, Middlesex, UK: Penguin, 1978).

  34. K. Kristiansen, “The Bronze Age Expansion of Indo-European Languages: An Archaeological Model,” in Becoming European: The Transformation of Third Millennium Northern and Western Europe, ed. Christopher Prescott and Håkon Glørstad (Oxford: Oxbow Books, 2011), 165–81.

  35. S. Rasmussen et al., “Early Divergent Strains of Yersinia pestis in Eurasia 5,000 Years Ago,” Cell 163 (2015): 571–82.