Modern humans are an anomaly in evolution, and the final key features occurred late in human evolution. Ultimate explanations for this evolutionary trajectory are best attained through synthetic studies that integrate genetics, biological anthropology, and archaeology, all resting firmly in the field of evolutionary anthropology. These fields of endeavor typically operate in relative isolation. This synthetic overview identifies the three pillars of human uniqueness: an evolved advanced cognition, hyperprosociality, and psychology for social learning. These properties are foundational for cumulative culture, the dominant adaptation of our species. Although the line evolved in the direction of advancing cognition, the evidence shows that only modern humans evolved extreme levels of prosociality and social learning; this review offers an explanation. These three traits were in place ∼200–100 ka and produced a creature capable of extraordinary social and technological structures, but one that was also empowered to make war in large groups with advanced weapons. The advance out of Africa and the annihilation of other hominin taxa, and many unprepared megafauna, were assured.


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Literature Cited

  1. Ames CJH, Riel-Salvatore J, Collins BR. 2013. Why we need an alternative approach to the study of modern human behaviour. Can. J. Archaeol. 37:21–47 [Google Scholar]
  2. Andreasen NC. 2006. The Creative Brain New York: Penguin [Google Scholar]
  3. Argue D, Morwood MJ, Sutikna T, Jatmiko, Saptomo EW. 2009. Homo floresiensis: a cladistic analysis. J. Hum. Evol. 57:623–39 [Google Scholar]
  4. Balzeau A, Holloway RL, Grimaud-Hervé D. 2012. Variations and asymmetries in regional brain surface in the genus Homo. J. Hum. Evol. 62:696–706 [Google Scholar]
  5. Bar-Yosef O, Bordes J-G. 2010. Who were the makers of the Châtelperronian culture?. J. Hum. Evol. 59:586–93 [Google Scholar]
  6. Bar-Yosef Mayer DE, Vandermeersch B, Bar-Yosef O. 2009. Shells and ochre in Middle Paleolithic Qafzeh Cave, Israel: indications for modern behavior. J. Hum. Evol. 56:307–14 [Google Scholar]
  7. Barham LS. 1998. Possible early pigment use in South-Central Africa. Curr. Anthropol. 39:703–10 [Google Scholar]
  8. Barham LS. 2002. Systematic pigment use in the Middle Pleistocene of South-Central Africa. Curr. Anthropol. 43:181–90 [Google Scholar]
  9. Barham LS. 2012. Clarifying some fundamental errors in Herries' “Chronological Perspective on the Acheulian and Its Transition to the Middle Stone Age in Southern Africa: The Question of the Fauresmith” (2011). Int. J. Evol. Biol. 2012:230156 [Google Scholar]
  10. Barton NH, Hewitt GM. 1989. Adaptation, speciation and hybrid zones. Nature 341:497–503 [Google Scholar]
  11. Berger TD, Trinkaus E. 1995. Patterns of trauma among the Neandertals. J. Archaeol. Sci. 22:841–52 [Google Scholar]
  12. Botha R. 2010. On the soundness of inferring modern language from symbolic behaviour. Cambridge Archaeol. J. 20:345–56 [Google Scholar]
  13. Bouzouggar A, Barton N, Vanhaeren M, d'Errico F, Collcutt S. et al. 2007. 82,000-year-old shell beads from North Africa and implications for the origins of modern human behavior. PNAS 104:9964–69 [Google Scholar]
  14. Bowles S. 2009. Did warfare among ancestral hunter-gatherers affect the evolution of human social behaviors?. Science 324:1293–98 [Google Scholar]
  15. Bowles S, Gintis H. 2011. A Cooperative Species: Human Reciprocity and Its Evolution Princeton, NJ: Princeton Univ. Press [Google Scholar]
  16. Boyd R, Richerson PJ, Henrich J. 2011. The cultural niche: why social learning is essential for human adaptation. PNAS 108:10918–25 [Google Scholar]
  17. Brown KS, Marean CW, Herries AIR, Jacobs Z, Tribolo C. et al. 2009. Fire as an engineering tool of early modern humans. Science 325:859–62 [Google Scholar]
  18. Brown KS, Marean CW, Jacobs Z, Schoville BJ, Oestmo S. et al. 2012. An early and enduring advanced technology originating 71,000 years ago in South Africa. Nature 491:590–93 [Google Scholar]
  19. Bruner E. 2010. Morphological differences in the parietal lobes within the human genus: a neurofunctional perspective. Curr. Anthropol. 51:S77–88 [Google Scholar]
  20. Caron F, d'Errico F, Del Moral P, Santos F, Zilhão J. et al. 2011. The reality of Neandertal symbolic behavior at the Grotte du Renne, Arcy-sur-Cure, France. PLOS ONE 6:e21545 [Google Scholar]
  21. Chan EKF, Hardie R-A, Petersen DC, Beeson K, Bornman RM. et al. 2015. Revised timeline and distribution of the earliest diverged human maternal lineages in Southern Africa. PLOS ONE 10:e0121223 [Google Scholar]
  22. Charrié-Duhaut A, Porraz G, Cartwright CR, Igreja M, Connan J. et al. 2013. First molecular identification of a hafting adhesive in the Late Howiesons Poort at Diepkloof Rock Shelter (Western Cape, South Africa). J. Archaeol. Sci. 40:3506–18 [Google Scholar]
  23. Churchill SE, Franciscus RG, Kean-Peraza HA, Daniel JA, Warren BR. 2009. Shanidar 3 Neandertal rib puncture wound and paleolithic weaponry. J. Hum. Evol. 57:163–78 [Google Scholar]
  24. Clark JD. 1988. The Middle Stone Age of East Africa and the beginnings of regional identity. J. World Prehistory 2:235–305 [Google Scholar]
  25. Clark JD, Beyene Y, WoldeGabriel G, Hart WK, Renne PR. et al. 2003. Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature 423:747–52 [Google Scholar]
  26. Clark JD, Williamson KD, Michels JW, Marean CW. 1984. A Middle Stone Age occupation site at Porc Epic cave, Dire Dawa (east-central Ethiopia). Afr. Archaeol. Rev. 2:37–71 [Google Scholar]
  27. Clutton-Brock T. 2002. Breeding together: kin selection and mutualism in cooperative vertebrates. Science 296:69–72 [Google Scholar]
  28. Coolidge FL, Wynn T. 2009. The Rise of Homo sapiens: The Evolution of Modern Thinking Oxford, UK: Wiley-Blackwell [Google Scholar]
  29. Cortés-Sánchez M, Morales-Muñiz A, Simón-Vallejo MD, Lozano-Francisco MC, Vera-Peláez JL. et al. 2011. Earliest known use of marine resources by Neanderthals. PLOS ONE 6:e24026 [Google Scholar]
  30. Crabtree DE, Butler BR. 1964. Notes on experiments in flint knapping: 1 Heat treatment of silica materials. Tebiwa 7:1–6 [Google Scholar]
  31. d'Errico F. 2003. The invisible frontier. A multiple species model for the origin of behavioral modernity. Evol. Anthropol. 12:188–202 [Google Scholar]
  32. d'Errico F, Banks WE. 2013. Identifying mechanisms behind Middle Paleolithic and Middle Stone Age cultural trajectories. Curr. Anthropol. 54:S371–87 [Google Scholar]
  33. d'Errico F, Henshilwood CS. 2007. Additional evidence for bone technology in the southern African Middle Stone Age. J. Hum. Evol. 52:142–63 [Google Scholar]
  34. d'Errico F, Stringer CB. 2011. Evolution, revolution or saltation scenario for the emergence of modern cultures?. Philos. Trans. R. Soc. B 366:1060–69 [Google Scholar]
  35. Day MH. 1969. Omo human skeletal remains. Nature 222:1135–38 [Google Scholar]
  36. Ding Q, Hu Y, Xu S, Wang J, Jin J. 2013. Neanderthal introgression at Chromosome 3p21.31 was under positive natural selection in East Asians. Mol. Biol. Evol. 31:683–95 [Google Scholar]
  37. Endicott P, Ho SYW, Stringer C. 2010. Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern human origins. J. Hum. Evol. 59:87–95 [Google Scholar]
  38. EPICA Community Memb 2004. Eight glacial cycles from an Antarctic ice core. Nature 429:623–28 [Google Scholar]
  39. Fagundes NJR, Ray N, Beaumont M, Neuenschwander S, Salzano FM. et al. 2007. Statistical evaluation of alternative models of human evolution. PNAS 104:17614–19 [Google Scholar]
  40. Féblot-Augustins J. 1993. Mobility strategies in the Late Middle Palaeolithic of Central Europe and Western Europe: elements of stability and variability. J. Anthropol. Archaeol. 12:211–65 [Google Scholar]
  41. Fehr E, Fischbacher U. 2003. The nature of human altruism. Nature 425:785–91 [Google Scholar]
  42. Fehr E, Gächter S. 2002. Altruistic punishment in humans. Nature 415:137–40 [Google Scholar]
  43. Fisher SE, Scharff C. 2009. FOXP2 as a molecular window into speech and language. Trends Genet. 25:166–77 [Google Scholar]
  44. Fu Q, Li H, Moorjani P, Jay F, Slepchenko SM, Bondarev AA. et al. 2014. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature 514:445–49 [Google Scholar]
  45. Gamble C. 1994. Timewalkers: The Prehistory of Global Colonization Cambridge, MA: Harvard Univ. Press [Google Scholar]
  46. Godfrey-Smith DI, Ilani S. 2004. Past thermal history of goethite and hematite fragments from Qafzeh Cave deduced from thermal activation characteristics of the 110°C TL peak of enclosed quartz grains. Rev. Archéometrie 28:185–90 [Google Scholar]
  47. Goren-Inbar N, Alperson N, Kislev ME, Simchoni O, Melamed Y. et al. 2004. Evidence of hominin control of fire at Gesher Benot Yaàqov, Israel. Science 304:725–27 [Google Scholar]
  48. Green RE, Krause J, Briggs AW, Maricic T, Stenzel U. et al. 2010. A draft sequence of the Neandertal genome. Science 328:710–22 [Google Scholar]
  49. Hammer MF, Woerner AE, Mendez FL, Watkins JC, Wall JD. 2011. Genetic evidence for archaic admixture in Africa. PNAS 108:15123–28 [Google Scholar]
  50. Harvati K, Stringer C, Grün R, Aubert M, Allsworth-Jones P, Polorunso CA. 2011. The Later Stone Age calvaria from Iwo Eleru, Nigeria: morphology and chronology. PLOS ONE 6:e24024 [Google Scholar]
  51. Henn BM, Cavalli-Sforza LL, Feldman MW. 2012. The great human expansion. PNAS 109:17758–64 [Google Scholar]
  52. Henn BM, Gignoux CR, Jobin M, Granka JM, Macpherson JM. et al. 2011. Hunter-gatherer genomic diversity suggests a southern African origin for modern humans. PNAS 108:5154–62 [Google Scholar]
  53. Henrich J. 2004. Cultural group selection, coevolutionary processes and large-scale cooperation. J. Econ. Behav. Organ. 53:3–35 [Google Scholar]
  54. Henrich J, Boyd R, Bowles S, Camerer C, Fehr E. et al. 2001. In search of Homo economicus: behavioral experiments in 15 small-scale societies. Am. Econ. Rev. 91:73–78 [Google Scholar]
  55. Henrich J, Ensminger J, McElreath R, Barr A, Barrett C. et al. 2010. Markets, religion, community size, and the evolution of fairness and punishment. Science 327:1480–84 [Google Scholar]
  56. Henrich J, Henrich N. 2006. Culture, evolution and the puzzle of human cooperation. Cogn. Syst. Res. 7:220–45 [Google Scholar]
  57. Henrich J, McElreath R. 2003. The evolution of cultural evolution. Evol. Anthropol. 12:123–35 [Google Scholar]
  58. Henshilwood C, d'Errico F, Vanhaeren M, van Niekerk K, Jacobs Z. 2004. Middle Stone Age shell beads from South Africa. Science 304:404–4 [Google Scholar]
  59. Henshilwood CS, d'Errico F, Marean CW, Milo RG, Yates R. 2001. An early bone tool industry from the Middle Stone Age at Blombos Cave, South Africa: implications for the origins of modern human behaviour, symbolism and language. J. Hum. Evol. 41:631–78 [Google Scholar]
  60. Henshilwood CS, d'Errico F, van Niekerk KL, Coquinot Y, Jacobs Z. et al. 2011. A 100,000-year-old ochre-processing workshop at Blombos Cave, South Africa. Science 334:219–22 [Google Scholar]
  61. Henshilwood CS, d'Errico F, Watts I. 2009. Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa. J. Hum. Evol. 57:27–47 [Google Scholar]
  62. Henshilwood CS, D'Errico F, Yates R, Jacobs Z, Tribolo C. et al. 2002. Emergence of modern human behavior: Middle Stone Age engravings from South Africa. Science 295:1278–80 [Google Scholar]
  63. Henshilwood CS, Dubreuil B. 2009. Reading the artefacts: gleaning language skills from the Middle Stone Age in Southern Africa. The Cradle of Language R Botha, C Knight 41–61 Oxford, UK: Oxford Univ. Press [Google Scholar]
  64. Henshilwood CS, Dubreuil B. 2011. The Still Bay and Howiesons Poort, 77–59 ka: symbolic material culture and the evolution of the mind during the African Middle Stone Age. Curr. Anthropol. 52:361–400 [Google Scholar]
  65. Henshilwood CS, Marean CW. 2003. The origin of modern human behavior: a review and critique of models and test implications. Curr. Anthropol. 44:627–51 [Google Scholar]
  66. Herries AIR. 2011. A chronological perspective on the Acheulian and its transition to the Middle Stone Age in southern Africa: the question of the Fauresmith. Int. J. Evol. Biol. 2011:961401 [Google Scholar]
  67. Hershkovitz I, Marder O, Ayalon A, Bar-Matthews M, Yasur G. et al. 2015. Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans. Nature 520:216–19 [Google Scholar]
  68. Hewitt G. 2011. Quaternary phylogeography: the roots of hybrid zones. Genetica 139:617–38 [Google Scholar]
  69. Higham T. 2011. European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies. Antiquity 85:235–49 [Google Scholar]
  70. Higham T, Douka K, Wood R, Ramsey CB, Brock F. et al. 2014. The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512:306–9 [Google Scholar]
  71. Hill K. 2002. Altruistic cooperation during foraging by the Ache, and the evolved human predisposition to cooperate. Hum. Nat. 13:105–28 [Google Scholar]
  72. Hill K, Barton CM, Hurtado AM. 2009. The emergence of human uniqueness: characters underlying behavioral modernity. Evol. Anthropol. 18:187–200 [Google Scholar]
  73. Hill K, Hurtado AM. 2009. Cooperative breeding in South American hunter-gatherers. Proc. R. Soc. B 276:3863–70 [Google Scholar]
  74. Holloway RL, Schwartz JH, Tattersall I. 2004. The Human Fossil Record, Brain Endocasts—The Paleoneurological Evidence New Jersey: Wiley [Google Scholar]
  75. House BR, Silk JB, Henrich J, Barrett HC, Scelza BA. et al. 2013. Ontogeny of prosocial behavior across diverse societies. PNAS 110:14586–91 [Google Scholar]
  76. Hovers E, Ilani S, Bar-Yosef O, Vandermeersch B. 2003. An early case of color symbolism. Curr. Anthropol. 44:491–522 [Google Scholar]
  77. Hrdy SB. 2011. Mothers and Others: The Evolutionary Origins of Mutual Understanding Cambridge, UK: Belknap [Google Scholar]
  78. Hublin JJ. 1992. Recent human evolution in northwestern Africa. Philos. Trans. R. Soc. B 337:185–91 [Google Scholar]
  79. Hublin JJ. 2009. The origin of Neandertals. PNAS 106:16022–27 [Google Scholar]
  80. Huerta-Sánchez E, Jin X, Asan, Bianba Z, Peter BM. et al. 2014. Altitude adaptation in Tibetans caused by introgression of Denisovan-like DNA. Nature 512:194–97 [Google Scholar]
  81. Jacobs Z, Roberts RG. 2007. Advances in optically stimulated luminescence dating of individual grains of quartz from archeological deposits. Evol. Anthropol. 16:210–23 [Google Scholar]
  82. Jerardino A, Marean CW. 2010. Shellfish gathering, marine paleoecology and modern human behavior: perspectives from cave PP13B, Pinnacle Point, South Africa. J. Hum. Evol. 59:412–24 [Google Scholar]
  83. Kaplan H, Hill K, Lancaster J, Hurtado AM. 2000. A theory of human life history evolution: diet, intelligence, and longevity. Evol. Anthropol. 9:156–85 [Google Scholar]
  84. Kim HL, Ratan A, Perry GH, Montenegro A, Miller W, Schuster SC. 2014. Khoisan hunter-gatherers have been the largest population throughout most of modern-human demographic history. Nat. Commun. 5:5692 [Google Scholar]
  85. Klein RG. 2000. Archeology and the evolution of human behavior. Evol. Anthropol. 9:17–36 [Google Scholar]
  86. Koller J, Baumer U, Mania D. 2001. High-tech in the Middle Palaeolithic: Neandertal-manufactured pitch identified. Eur. J. Archaeol. 4:385–97 [Google Scholar]
  87. Kuhn SL. 2013. Roots of the Middle Paleolithic in Eurasia. Curr. Anthropol. 54:S255–68 [Google Scholar]
  88. Lachance J, Vernot B, Elbers CC, Ferwerda B, Froment A. et al. 2012. Evolutionary history and adaptation from high-coverage whole-genome sequences of diverse African hunter-gatherers. Cell 150:457–69 [Google Scholar]
  89. Lahr MM, Foley RA. 1998. Towards a theory of modern human origins: geography, demography, and diversity in recent human evolution. Yearb. Phys. Anthropol. 41:137–76 [Google Scholar]
  90. Larick R. 1985. Spears, style, and time among Maa-speaking pastoralists. J. Anthropol. Archaeol. 4:206–20 [Google Scholar]
  91. Lieberman P. 2007. The evolution of human speech: its anatomical and neural bases. Curr. Anthropol. 48:39–66 [Google Scholar]
  92. Lombard M, Haidle MN. 2012. Thinking a bow-and-arrow set: cognitive implications of Middle Stone Age bow and stone-tipped arrow technology. Cambridge Archaeol. J. 22:237–64 [Google Scholar]
  93. Mackay A, Welz A. 2008. Engraved ochre from a Middle Stone Age context at Klein Kliphuis in the Western Cape of South Africa. J. Archaeol. Sci. 35:1521–32 [Google Scholar]
  94. Marean CW. 2010. Pinnacle Point Cave 13B (Western Cape Province, South Africa) in context: the Cape Floral kingdom, shellfish, and modern human origins. J. Hum. Evol. 59:425–43 [Google Scholar]
  95. Marean CW. 2011. Coastal South Africa and the co-evolution of the modern human lineage and coastal adaptations. Trekking the Shore: Changing Coastlines and the Antiquity of Coastal Settlement N Bicho, JA Haws, LG Davis 421–40 New York: Springer [Google Scholar]
  96. Marean CW. 2014. The origins and significance of coastal resource use in Africa and Western Eurasia. J. Hum. Evol. 77:17–40 [Google Scholar]
  97. Marean CW, Assefa Z. 1999. Zooarchaeological evidence for the faunal exploitation behavior of Neandertals and early modern humans. Evol. Anthropol. 8:22–37 [Google Scholar]
  98. Marean CW, Assefa Z. 2005. The Middle and Upper Pleistocene African record for the biological and behavioral origins of modern humans. African Archaeology AB Stahl 93–129 New York: Blackwell [Google Scholar]
  99. Marean CW, Bar-Matthews M, Bernatchez J, Fisher E, Goldberg P. et al. 2007. Early human use of marine resources and pigment in South Africa during the Middle Pleistocene. Nature 449:905–8 [Google Scholar]
  100. Marean CW, Cawthra HC, Cowling RM, Esler KJ, Fisher E. et al. 2014. Stone Age people in a changing South African Greater Cape Floristic Region. Fynbos: Ecology, Evolution, and Conservation of a Megadiverse Region N Allsopp, JF Colville, GA Verboom 164–99 Oxford, UK: Oxford Univ. Press [Google Scholar]
  101. Mazza PPA, Martini F, Sala B, Magi M, Colombini MP. et al. 2006. A new Palaeolithic discovery: tar-hafted stone tools in a European Mid-Pleistocene bone-bearing bed. J. Archaeol. Sci. 33:1310–18 [Google Scholar]
  102. McBrearty S, Brooks AS. 2000. The revolution that wasn't: a new interpretation of the origin of modern human behavior. J. Hum. Evol. 39:453–563 [Google Scholar]
  103. McDougall I, Brown FH, Fleagle JG. 2005. Stratigraphic placement and age of modern humans from Kibish, Ethiopia. Nature 433:733–36 [Google Scholar]
  104. McDougall I, Brown FH, Fleagle JG. 2008. Sapropels and the age of hominins Omo I and II, Kibish, Ethiopia. J. Hum. Evol. 55:409–20 [Google Scholar]
  105. McGuigan N, Makinson J, Whiten A. 2011. From over-imitation to super-copying: Adults imitate causally irrelevant aspects of tool use with higher fidelity than young children. Br. J. Psychol. 102:1–18 [Google Scholar]
  106. Mellars P, Stringer C. 1989. The Human Revolution: Behavioural and Biological Perspectives on the Origins of Modern Humans Edinburgh: Edinb. Univ. Press [Google Scholar]
  107. Mellars P, Stringer C, Bar-Yosef O, Boyle K. 2007. Rethinking the Human Revolution Cambridge, UK: MacDonald Inst. Archaeol. Res. [Google Scholar]
  108. Mendez FL, Krahn T, Schrack B, Krahn AM, Veeramah KR. et al. 2013. An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree. Am. J. Hum. Genet. 92:454–59 [Google Scholar]
  109. Merrick HV, Brown FH. 1984. Obsidian sources and patterns of source utilization in Kenya and northern Tanzania: some initial findings. Afr. Archaeol. Rev. 2:129–52 [Google Scholar]
  110. Merrick HV, Brown FH, Nash WP. 1994. Use and movement of obsidian in the Early and Middle Stone Ages of Kenya and Northern Tanzania. Society, Culture, and Technology in Africa ST Childs 29–44 Philadelphia: Univ. Pa. Mus. Archaeol. Anthropol. [Google Scholar]
  111. Meyer M, Fu Q, Aximu-Petri A, Glocke I, Nickel B. et al. 2014. A mitochondrial genome sequence of a hominin from Sima de los Huesos. Nature 505:403–6 [Google Scholar]
  112. Nash DJ, Coulson S, Staurset S, Ullyott JS, Babutsi M. et al. 2013. Provenancing of silcrete raw materials indicates long-distance transport to Tsodilo Hills, Botswana, during the Middle Stone Age. J. Hum. Evol. 64:280–88 [Google Scholar]
  113. Negash A, Brown F, Nash B. 2011. Varieties and sources of artefactual obsidian in the Middle Stone Age of the Middle Awash, Ethiopia. Archaeometry 53:661–73 [Google Scholar]
  114. Negash A, Shackley MS. 2006. Geochemical provenance of obsidian artefacts from the MSA site of Porc Epic, Ethiopia. Archaeometry 48:1–12 [Google Scholar]
  115. Nielsen M, Mushin I, Tomaselli K, Whiten A. 2014. Where culture takes hold: “overimitation” and its flexible deployment in Western, Aboriginal, and Bushmen children. Child Dev. 85:2169–84 [Google Scholar]
  116. Nowell A. 2010. Defining behavioral modernity in the context of Neandertal and anatomically modern human populations. Annu. Rev. Anthropol. 39:437–52 [Google Scholar]
  117. Pearce E, Stringer C, Dunbar RIM. 2013. New insights into differences in brain organization between Neanderthals and anatomically modern humans. Proc. R. Soc. B 280:20130168 [Google Scholar]
  118. Perreault C, Mathew S. 2012. Dating the origin of language using phonemic diversity. PLOS ONE 7:e35289 [Google Scholar]
  119. Pickrell JK, Patterson N, Barbieri C, Berthold F, Gerlach L. et al. 2012. The genetic prehistory of southern Africa. Nat. Commun. 3:1143 [Google Scholar]
  120. Pickrell JK, Reich D. 2014. Toward a new history and geography of human genes informed by ancient DNA. Trends Genet. 30:377–89 [Google Scholar]
  121. Pinker S. 2010. The cognitive niche: coevolution of intelligence, sociality, and language. PNAS 107:8993–99 [Google Scholar]
  122. Ponce de León MS, Golovanova L, Doronichev V, Romanova G, Akazawa T. et al. 2008. Neanderthal brain size at birth provides insights into the evolution of human life history. PNAS 105:13764–68 [Google Scholar]
  123. Powell A, Shennan S, Thomas MG. 2009. Late Pleistocene demography and the appearance of modern human behavior. Science 324:1298–301 [Google Scholar]
  124. Poznik GD, Henn BM, Yee M-C, Sliwerska E, Euskirchen GM. et al. 2013. Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females. Science 341:562–65 [Google Scholar]
  125. Prüfer K, Racimo F, Patterson N, Jay F, Sankararaman S. et al. 2014. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature 505:43–49 [Google Scholar]
  126. Rasmussen M, Guo X, Wang Y, Lohmueller KE, Rasmussen S. et al. 2011. An Aboriginal Australian genome reveals separate human dispersals into Asia. Science 334:94–98 [Google Scholar]
  127. Reich D, Patterson N, Kircher M, Delfin F, Nandineni MR. et al. 2011. Denisova admixture and the first modern human dispersals into Southeast Asia and Oceania. Am. J. Hum. Genet. 89:516–28 [Google Scholar]
  128. Rhodes JA, Churchill SE. 2009. Throwing in the Middle and Upper Paleolithic: inferences from an analysis of humeral retroversion. J. Hum. Evol. 56:1–10 [Google Scholar]
  129. Richerson PJ, Boyd R. 2000. Built for speed: pleistocene climate variation and the origin of human culture. Perspect. Ethol. 13:1–45 [Google Scholar]
  130. Richerson PJ, Boyd R. 2005. Not by Genes Alone Chicago: Univ. Chicago Press [Google Scholar]
  131. Rightmire GP. 1979. Implications of Border Cave skeletal remains for Later Pleistocene human evolution. Curr. Anthropol. 20:23–35 [Google Scholar]
  132. Roebroeks W, Sier MJ, Nielsen TK, De Loecker D, Parés JM. et al. 2012. Use of red ochre by early Neandertals. PNAS 109:1889–94 [Google Scholar]
  133. Salomon H, Vignaud C, Coquinot Y, Beck L, Stringer C. et al. 2012. Selection and heating of colouring materials in the Mousterian level of Es-Skhul (c. 100 000 years BP, Mount Carmel, Israel). Archaeometry 54:698–722 [Google Scholar]
  134. Sankararaman S, Mallick S, Dannemann M, Prüfer K, Kelso J. et al. 2014. The genomic landscape of Neanderthal ancestry in present-day humans. Nature 507:354–57 [Google Scholar]
  135. Scally A, Durbin R. 2012. Revising the human mutation rate: implications for understanding human evolution. Nat. Rev. Genet. 13:745–53 [Google Scholar]
  136. Schlebusch C, Lombard M, Soodyall H. 2013. MtDNA control region variation affirms diversity and deep sub-structure in populations from southern Africa. BMC Evol. Biol. 13:56 [Google Scholar]
  137. Schlebusch CM, Skoglund P, Sjöin P, Gattepaille LM, Hernandez D. et al. 2012. Genomic variation in seven Khoe-San groups reveals adaptation and complex African history. Science 338:374–79 [Google Scholar]
  138. Schwartz GT. 2012. Growth, development, and life history throughout the evolution of Homo. Curr. Anthropol. 53:S395–408 [Google Scholar]
  139. Shea JJ. 2006. The origins of lithic projectile point technology: evidence from Africa, the Levant, and Europe. J. Archaeol. Sci. 33:823–46 [Google Scholar]
  140. Shea JJ. 2011. Homo sapiens Is as Homo sapiens Was. Curr. Anthropol. 52:1–35 [Google Scholar]
  141. Shultz S, Nelson E, Dunbar RIM. 2012. Hominin cognitive evolution: identifying patterns and processes in the fossil and archaeological record. Philos. Trans. R. Soc. B 367:2130–40 [Google Scholar]
  142. Singer R, Wymer J. 1982. The Middle Stone Age at Klasies River Mouth in South Africa Chicago: Univ. Chicago Press [Google Scholar]
  143. Smith EA. 2013. Agency and adaptation: new directions in evolutionary anthropology. Annu. Rev. Anthropol. 42:103–20 [Google Scholar]
  144. Smith TM, Tafforeau P, Reid DJ, Grün R, Eggins S. et al. 2007. Earliest evidence of modern human life history in North African early Homo sapiens. PNAS 104:6128–33 [Google Scholar]
  145. Smith TM, Tafforeau P, Reid DJ, Pouech J, Lazzari V. et al. 2010. Dental evidence for ontogenetic differences between modern humans and Neanderthals. PNAS 107:20923–28 [Google Scholar]
  146. Soressi M, Rendu W, Texier J-P, Claud L, Daulny L. et al. 2008. Pech-de-l'Azé I (Dordogne, France): nouveau regard sur un gisement moustérien de tradition acheuléenne connu depuis le XIXe siècle. Bull. Soc. Préhist. Fr. 47:95–132 [Google Scholar]
  147. Stringer C. 2012. The status of Homo heidelbergensis (Schoetensack 1908). Evol. Anthropol. 21:101–7 [Google Scholar]
  148. Stringer CB, Finlayson JC, Barton RNE, Fernández-Jalvo Y, Cáceres I. et al. 2008. Neanderthal exploitation of marine mammals in Gibraltar. PNAS 105:14319–24 [Google Scholar]
  149. Tennie C, Call J, Tomasello M. 2009. Ratcheting up the ratchet: on the evolution of cumulative culture. Philos. Trans. R. Soc. B 364:2405–15 [Google Scholar]
  150. Texier P-J, Porraz G, Parkington J, Rigaud J-P, Poggenpoel C. et al. 2010. A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa. PNAS 107:6180–85 [Google Scholar]
  151. Tishkoff S, Reed FA, Friedlaender FR, Ehret C, Ranciaro A. et al. 2009. The genetic structure and history of Africans and African Americans. Science 324:1035–44 [Google Scholar]
  152. Tomasello M. 1999. The Cultural Origins of Human Cognition Cambridge, MA: Harvard Univ. Press [Google Scholar]
  153. Tomasello M. 2008. Origins of Human Communication Cambridge, MA: MIT Press [Google Scholar]
  154. Tomasello M, Kruger AC, Ratner HH. 1993. Cultural learning. Behav. Brain Sci. 16:495–511 [Google Scholar]
  155. Trinkaus E. 1983. The Shanidar Neandertals New York: Academic [Google Scholar]
  156. Trinkaus E. 1995. Neanderthal mortality patterns. J. Archaeol. Sci. 22:121–42 [Google Scholar]
  157. Underhill PA, Kivisild T. 2007. Use of Y chromosome and mitochondrial DNA population structure in tracing human migrations. Annu. Rev. Genet. 41:539–64 [Google Scholar]
  158. Vaks A, Bar-Matthews M, Ayalon A, Matthews A, Halicz L, Frumkin A. 2007. Desert speleothems reveal climatic window for African exodus of early modern humans. Geology 35:831–34 [Google Scholar]
  159. Valladas H, Reyss J-L, Vallardas G, Bar-Yosef O, Vandermeersch B. 1988. Thermoluminescence dating of Mousterian “Proto-Cro-Magnon” remains from Israel and the origin of modern man. Nature 331:614–16 [Google Scholar]
  160. Vanhaeren M, d'Errico F. 2006. Aurignacian ethno-linguistic geography of Europe revealed by personal ornaments. J. Archaeol. Sci. 33:1105–28 [Google Scholar]
  161. Wadley L. 2003. How some archaeologists recognize culturally modern behaviour. South Afr. J. Sci. 99:247–50 [Google Scholar]
  162. Wadley L, Williamson B, Lombard M. 2004. Ochre as a hafting material of Middle Stone Age stone tools: the evidence from Sibudu Cave, Kwazulu-Natal, South Africa. Antiquity 78:661–75 [Google Scholar]
  163. Watts I. 2002. Ochre in the Middle Stone Age of southern Africa: ritualized display or hide preservative?. S. Afr. Archaeol. Bull. 57:1–14 [Google Scholar]
  164. Watts I. 2010. The pigments from Pinnacle Point Cave 13B, Western Cape, South Africa. J. Hum. Evol. 59:392–411 [Google Scholar]
  165. White TD, Asfaw B, DeGusta D, Gilbert H, Richards GD. et al. 2003. Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature 423:742–47 [Google Scholar]
  166. Wiessner P. 1983. Style and social information in Kalahari San projectile points. Am. Antiq. 48:253–76 [Google Scholar]
  167. Zilhão J, Angelucci D, Badal-Garcia E, d'Errico F, Daniel F. et al. 2010. Symbolic use of marine shells and mineral pigments by Iberian Neandertals. PNAS 107:1023–28 [Google Scholar]

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