Rethinking the dispersal of Homo sapiens out of Africa
Huw S. Groucutt
Search for more papers by this authorMichael D. Petraglia
Search for more papers by this authorGeoff Bailey
Search for more papers by this authorEleanor M. L. Scerri
Search for more papers by this authorAsh Parton
Search for more papers by this authorLaine Clark-Balzan
Search for more papers by this authorRichard P. Jennings
Search for more papers by this authorLaura Lewis
Search for more papers by this authorJames Blinkhorn
Search for more papers by this authorNick A. Drake
Search for more papers by this authorPaul S. Breeze
Search for more papers by this authorRobyn H. Inglis
Search for more papers by this authorMaud H. Devès
Search for more papers by this authorMatthew Meredith-Williams
Search for more papers by this authorNicole Boivin
Search for more papers by this authorMark G. Thomas
Search for more papers by this authorAylwyn Scally
Search for more papers by this authorHuw S. Groucutt
Search for more papers by this authorMichael D. Petraglia
Search for more papers by this authorGeoff Bailey
Search for more papers by this authorEleanor M. L. Scerri
Search for more papers by this authorAsh Parton
Search for more papers by this authorLaine Clark-Balzan
Search for more papers by this authorRichard P. Jennings
Search for more papers by this authorLaura Lewis
Search for more papers by this authorJames Blinkhorn
Search for more papers by this authorNick A. Drake
Search for more papers by this authorPaul S. Breeze
Search for more papers by this authorRobyn H. Inglis
Search for more papers by this authorMaud H. Devès
Search for more papers by this authorMatthew Meredith-Williams
Search for more papers by this authorNicole Boivin
Search for more papers by this authorMark G. Thomas
Search for more papers by this authorAylwyn Scally
Search for more papers by this authorHuw S. Groucutt, Eleanor M. L. Scerri, Michael D. Petraglia, Ash Parton, Laine Clark-Balzan, Richard P. Jennings, Laura Lewis, Nicole Boivin School of Archaeology, Research Laboratory for Archaeology and the History of Art, University of Oxford, Oxford, OX1 3QY, United Kingdom. Corresponding author: Huw Groucutt. Email: [email protected]
Geoff Bailey, Robyn H. Inglis, Maud H. Devès, Matthew Meredith-Williams Department of Archaeology, University of York, York, YO1 7EP, United Kingdom
James Blinkhorn McDonald Institute for Archaeological Research, University of Cambridge, Downing Street, Cambridge CB2 3DZ
Nick A. Drake, Paul S. Breeze Department of Geography, King's College London, WC2R 2LS, United Kingdom
Maud H. Devès Laboratoire Tectonique, Institut de Physique du Globe, Paris, 75252, France
Mark G. Thomas Research Department of Genetics, Evolution and Environment, University College London, London, WC1E 6BT, United Kingdom
Aylwyn Scally Department of Genetics, University of Cambridge, Cambridge, CB2 3EH, United Kingdom
Abstract
Current fossil, genetic, and archeological data indicate that Homo sapiens originated in Africa in the late Middle Pleistocene. By the end of the Late Pleistocene, our species was distributed across every continent except Antarctica, setting the foundations for the subsequent demographic and cultural changes of the Holocene. The intervening processes remain intensely debated and a key theme in hominin evolutionary studies. We review archeological, fossil, environmental, and genetic data to evaluate the current state of knowledge on the dispersal of Homo sapiens out of Africa. The emerging picture of the dispersal process suggests dynamic behavioral variability, complex interactions between populations, and an intricate genetic and cultural legacy. This evolutionary and historical complexity challenges simple narratives and suggests that hybrid models and the testing of explicit hypotheses are required to understand the expansion of Homo sapiens into Eurasia.
REFERENCES
- 1 Willoughby PR. 2007. The evolution of modern humans in Africa; a comprehensive guide. New York: Altimira.
- 2 Petraglia MD, Haslam M, Fuller DQ, et al. 2010. Out of Africa: new hypotheses and evidence for the dispersal of Homo sapiens along the Indian Ocean rim. Ann Hum Biol 37: 288–311.
- 3 Stringer C. 2011. The origin of our species. London: Penguin.
- 4 Stringer C. 2014. Why we aren't all multiregionalists now. Trends Ecol Evol 29: 248–251.
- 5 McCown TD, Keith A. 1939. The Stone Age of Mt. Carmel, vol 2. The fossil human remains from the Levalloiso-Mousterian. Oxford: Clarendon Press.
- 6 Field H. 1932. The cradle of Homo sapiens. J Archaeol 36: 426–430.
- 7 Thorne AG, Wolpoff MH. 2003. The multiregional evolution of humans (revised paper). Sci Am 13: 46–53.
- 8 Bräuer G. 1992. Africa's place in the evolution of Homo sapiens. In: G Bräuer, F Smith, editors. Continuity or replacement? Controversies in Homo sapiens evolution. Rotterdam: Balkema. p 83–98.
- 9 Green RE, Krause J, Briggs AW, et al. 2010. A draft sequence of the Neandertal genome. Science 328: 710–722.
- 10 Reich D, Patterson N, Kircher M, et al. 2011. Denisova admixture and the first modern human dispersals into southeast Asia and Oceania. Am J Hum Genet 89: 516–528.
- 11 Smith F. 1992. The role of continuity in modern human origins. In: G Bräuer, F Smith, editors. Continuity or replacement? Controversies in Homo sapiens evolution. Rotterdam: Balkema. p 145–156.
- 12 Trinkaus E. 2005. Early modern humans. Ann Rev Anthropol 34: 207–230.
- 13 Lahr MM, Foley RA. 1998. Towards a theory of modern human origins: geography, demography and diversity in recent human evolution. Yearbk Phys Anthropol 41: 137–176.
- 14 Klein RG. 2009. The human career: human biological and cultural origins. Chicago: Chicago University Press.
10.7208/chicago/9780226027524.001.0001 Google Scholar
- 15 Shea JJ. 2011. The archaeology of an illusion: the Middle-Upper Paleolithic transition in the Levant. In: JM Tenesor, R Jagher, R Otte, editors. The Lower and Middle Paleolithic in the Middle East and neighbouring regions. ERAUL 126. Liége: Université de Liége. p 169–182.
- 16 Oppenheimer S. 2012. A single southern exit of modern humans from Africa: before or after Toba? Quat Int 258: 88–99.
- 17 Mellars P, Gori KC, Carr M, et al. 2013. Genetic and archaeological perspectives on the initial modern human colonization of southern Asia. Proc Natl Acad Sci USA 110: 10699–10704.
- 18 Reyes-Centeno H, Ghirotto S, Détroit F, et al. 2014. Genomic and cranial phenotype data support multiple modern human dispersals from Africa and a southern route into Asia. Proc Natl Acad Sci USA 111: 7248–7253.
- 19 Armitage SJ, Jasim SA, Marks AE, et al. 2011. The southern route “out of Africa”: evidence for an early expansion of modern humans into Arabia. Science 331: 453–456.
- 20 Usik VI, Rose JI, Hilbert YH, et al. 2013. Nubian Complex reduction strategies in Dhofar, southern Oman. Quat Int 300: 244–266.
- 21 Gunz P, Bookstein FL, Mitteroecker P, et al. 2009. Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario. Proc Natl Acad Sci USA 106: 6094–6098.
- 22 Douka K. 2013. Exploring “the great wilderness of prehistory”: the chronology of the Middle to the Upper Palaeolithic transition in the Northern Levant. Mitt Gessell Urgeschichte 22: 11–40.
- 23 Hershkovitz I, Marder O, Ayalon A. 2015. Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans. Nature 520: 216–219.
- 24 Shea JJ. 2003. The Middle Paleolithic of the East Mediterranean Levant. J World Prehist 17: 313–394.
- 25 Dennell R. 2014. Smoke and mirrors: the fossil record for Homo sapiens between Arabia and Australia. In: R Dennell, M Porr, editors. Southern Asia, Australia and the search for human origins. Cambridge: Cambridge University Press. p 33–50.
10.1017/CBO9781139084741.004 Google Scholar
- 26 Groucutt HS, Petraglia MD. 2012. The prehistory of the Arabian Peninsula: deserts, dispersals and demography. Evol Anthropol 21: 113–125.
- 27 Boivin N, Fuller DQ, Dennell R, et al. 2013. Human dispersal across diverse environments of Asia during the Upper Pleistocene. Quat Int 300: 32–47.
- 28 Mijares SR, Detroit F, Piper P, et al. 2010. New evidence for a 67,000-year-old human presence at Callao Cave, Luzon, Philippines. J Hum Evol 59: 123–132.
- 29 Demeter F, Shackelford L, Westaway K, et al. 2015. Early modern humans and morphological variation in Southeast Asia: fossil evidence from Tam Pa Ling, Laos. PLoS ONE 10: e0121193.
- 30 Cann R, Stoneking M, Wilson A. 1987. Mitochondrial DNA and human evolution. Nature 325: 31–36.
- 31 Hammer MF. 1995. A recent common ancestry for human Y chromosome. Nature 378: 376–378.
- 32 Mountain JL, Jin AA, Bowcock AM, et al. 1993. Evolution of modern humans: evidence from nuclear DNA polymorphisms. In: MJ Aitken, CB Stringer, PA Mellars, editors. The origin of modern humans and the impact of chronometric dating. Princeton University Press: Princeton.
- 33 Veeramah KR, Hammer MF. 2014. The impact of whole-genome sequencing on the reconstruction of human population history. Nat Rev Genet 15: 149–162.
- 34 Pickrell JK, Reich D. 2014. Toward a new history and geography of human genes informed by ancient DNA. Trends Genet 30: 377–389.
- 35 Seguin-Orlando A, Korneliussen TS, Sikora M. 2014. Genomic structure in Europeans dating back to least 36,000 years. Science 346: 1113–1118.
- 36 Soares P, Alshamali F, Pereira JB, et al. 2012. The expansion of mtDNA haplogroup L3 within and out of Africa. Mol Biol Evol 29: 915–927.
- 37 Pearson OM. 2013. Africa: the cradle of modern people. In: FH Smith, JCM Ahern, editors. The origins of modern humans: biology reconsidered. Hoboken: John Wiley & Sons. p 1–43.
- 38 Campbell MC, Tishkoff S. 2010. The evolution of human genetic and phenotypic variation in Africa. Curr Biol 20: R166–173.
- 39 Schiffels S, Durbin R. 2014. Inferring human population size and separation history from multiple genome sequences. Nat Genet 46: 919–925.
- 40 Scally A, Durbin R. 2012. Revising the human mutation rate: implications for understanding human evolution. Nat Rev Genet 13: 745–753.
- 41 Fu Q, Mittnik A, Johnson PLF, et al. 2013. A revised timescale for human evolution based on ancient mitochondrial genomes. Curr Biol 23: 553–559.
- 42 Ségurel L, Wyman J, Przeworski M. 2014. Determinants of mutation rate variables in the human germline. Ann Rev Genomics Hum Genet 15: 47–70.
- 43 Fu Q, Moorjani P, Jay F, et al. 2014. Genome sequencing of a 45,000-year-old modern human from western Siberia. Nature 514: 445–449.
- 44 Underwhill PA, Kivisild T. 2007. Use of Y chromosome and mitochondrial DNA population structure in tracing human migrations. Ann Rev Genet 41: 539–564.
- 45 Wei W, Ayub Q, Chen Y, et al. 2013. A calibrated human Y-chromosal phylogeny based on resequencing. Genome Res 23: 388–395.
- 46 Li H, Durbin R. 2011. Inference of human population history from individual whole-genome sequences. Nature 475: 493–496.
- 47 Pagani L, Kivisild T, Tarekegn A, et al. 2012. Ethiopian genetic diversity reveals linguistic stratification and complex influences on the Ethiopian gene pool. Am J Hum Genet 91: 83–96.
- 48 Gurdasani D, Carstensen T, Tekola-Ayele F, et al. 2014. The African genome variation project shapes medical genetics in Africa. Nature 517: 327–332.
- 49 Prüfer K, Racimo F, Patterson N, et al. 2014. The complete genome sequence of a Neandertal from the Altai Mountains. Nature 505: 43–49.
- 50 Currat M, Excoffier L. 2014. Strong reproductive isolation between humans and Neanderthals inferred from observed patterns of introgression. Proc Natl Acad Sci USA 108: 15129–15134.
- 51 Clarkson C, Smith M, Marwick B, et al. n.d. The archaeology, chronology and stratigraphy of Madjedbebe (Malakunanja II): a site in northern Australia with early occupation. J Hum Evol. In press. doi:10.1016/j.jhevol.2015.03.014.
10.1016/j.jhevol.2015.03.014 Google Scholar
- 52 Groucutt HS, Scerri EML, Lewis L, et al. n.d. Stone tool assemblages and models for the dispersal of Homo sapiens out of Africa. Quat Int. In press. doi:10.1016/j.quaint.2015.01.039
10.1016/j.quaint.2015.01.039 Google Scholar
- 53 Porraz G, Texier PJ, Archer W, et al. 2013. Technological successions in the Middle Stone Age sequence of Diepkloof Rock Shelter, Western Cape, South Africa. J Archaeol Sci 40: 3376–3400.
- 54 Lewis L, Perera N, Petraglia M. 2014. First technological comparison of Southern African Howiesons Poort and South Asian Microlithic industries: an exploration of inter-regional variability in microlithic assemblages. Quat Int 350: 7–25.
- 55 Brandt SA, Fisher EC, Hildebrand EA, et al. 2012. Early MIS 3 occupation of Mochena Borago rockshelter, southwest Ethiopian highlands: implications for Late Pleistocene archaeology, paleoenvironments and modern human dispersals. Quat Int 274: 38–54.
- 56 James HVA, Petraglia M. 2005. Modern human origins and the evolution of behavior in the later Pleistocene record of South Asia. Curr Anthropol 46(suppl): S4–S27.
- 57 Petraglia M, Clarkson C, Boivin N, et al. 2009. Population increase and environmental deterioration correspond with microlithic innovations in South Asia ca. 35,000 years ago. Proc Natl Acad Sci USA 106: 12261–12266.
- 58 Clarkson C, Petraglia MD, Korisettar R, et al. 2009. The oldest and longest enduring microlithic sequence in India: 35,000 years of modern human occupation at the Jwalapuram Locality 9 rockshelter. Antiquity 83: 326–348.
- 59 Perera N, Kourampas N, Simpson IA, et al. 2011. People of the ancient rainforest: Late Pleistocene foragers at the Batadomba-lena rockshelter, Sri Lanka. J Hum Evol 61: 254–269.
- 60 Blinkhorn, J. 2014. Late Middle Palaeolithic surface sites occurring on dated sediment formations in the Thar Desert. Quat Int 350: 95–104.
- 61 Deraniyagala SU. 1992. The prehistory of Sri Lanka: an ecological perspective. Colombo: Department of Archaeological Survey.
- 62 Seligman CG. 1921. The older Palaeolithic age in Egypt. J R Anthropol Inst GB Ireland 51: 115–153.
- 63 Shea JJ. 2008. The Middle Stone Age archaeology of the Lower Omo Valley Kibish formation: excavations, lithic assemblages, and inferred patterns of early Homo sapiens behavior. J Hum Evol 55: 448–485.
- 64 Wendorf F, Schild R, Close AE, et al. 1993. Egypt during the Last Interglacial: the Middle Paleolithic of Bir Tarfawi and Bir Sahara East. Plenum: New York.
10.1007/978-1-4615-2908-8 Google Scholar
- 65 Hovers E. 2009. The lithic assemblages of Qafzeh Cave. Oxford: Oxford University Press.
- 66 Clarkson C, Jones S, Harris C. 2012. Continuity and change in the lithic industries of the Jurreru Valley, India, before and after the Toba eruption. Quat Int 258: 165–179.
- 67 Blinkhorn J, Achyuthan H, Petraglia MD, et al. 2013. Middle Palaeolithic occupation in the Thar Desert during the Upper Pleistocene: the signature of a modern human exit out of Africa? Quat Sci Rev 77: 233–238.
- 68 Scerri EML, Drake NA, Jennings R, et al. 2014. Earliest evidence for the structure of Homo sapiens populations in Africa. Quat Sci Rev 101: 207–216.
- 69 Dennell RW. 2009. The Palaeolithic settlement of Asia. Cambridge: Cambridge University Press.
- 70 Rabett RJ. 2012. Human adaptation in the Asian Palaeolithic: hominin dispersal and behaviour during the Late Quaternary. Cambridge: Cambridge University Press.
10.1017/CBO9781139087582 Google Scholar
- 71 Vanhaeren M, d'Errico F, Stringer C, et al. 2006. Middle Paleolithic shell beads in Israel and Algeria. Science 312: 1785–1788.
- 72 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.
- 73 Shea JJ. 2011. Homo sapiens is as Homo sapiens was. Curr Anthropol 52: 1–35.
- 74 Powell A, Shennan S, Thomas M. 2009. Late Pleistocene demography and the appearance of modern human behavior. Science 324: 1298–1301.
- 75 Blome MA, Cohen AS, Tryon CA, et al. 2012. The environmental context for the origins of modern human diversity: a synthesis of regional variability in African climate 150,000–30,000 years ago. J Hum Evol 62: 563–592.
- 76 Drake NA, Blench MA, Armitage SJ, et al. 2011. Ancient watercourses and biogeography of the Sahara explain the peopling of the desert. Proc Natl Acad Sci USA 108: 458–462.
- 77 Lane CS, Chorn BT, Johnson TC. 2013. Ash from Toba supereruption in Lake Malawi shows no volcanic winter in East Africa at 75 ka. Proc Natl Acad Sci USA 110: 8025–8029.
- 78 Parton A, White TS, Parker AG, et al. n.d. Orbital-scale climate variability in Arabia as a potential motor for human dispersals. Quat Int. In press. doi:10.1016/j.quaint.2015.01.005.
10.1016/j.quaint.2015.01.005 Google Scholar
- 79 Walter RC, Buffler RT, Bruggemann JH, et al. 2000. Early human occupation of the Red Sea coast of Eritrea during the Last Interglacial. Nature 405: 65–9.
- 80 Erlandson JM. 2001. The archaeology of aquatic adaptations: paradigms for a new millennium. J Archaeol Res 9: 287–350.
- 81 Bailey GN, Milner NJ. 2002. Coastal hunters and gatherers and social evolution: marginal or central? Before Farming 3-4: 1–15.
10.3828/bfarm.2002.3-4.1 Google Scholar
- 82 Bailey GN, Flemming N. 2008. Archaeology of the continental shelf: marine resources, submerged landscapes and underwater archaeology. Quat Sci Rev 27: 2153–2165.
- 83 Marean CW. 2014. The origins and significance of coastal resource use in Africa and Western Eurasia. J Hum Evol 77: 17–40.
- 84 Bruggemann JH, Buffler RT, Guillaume MMM, et al. 2004. Stratigraphy, palaeoenvironments and model for the deposition of the Abdur Reef limestone: context for an important archaeological site from the last interglacial on the Red Sea. Palaeogeogr, Palaeoclimatol, Palaeoecol 20: 179–206.
- 85 Stringer CB, Finlayson JC, Barton RNE, et al. 2008. Neanderthal exploitation of marine mammals in Gibraltar. Proc Natl Acad Sci USA 108: 14319–14324.
- 86 Irwin G. 1992. The Prehistoric exploration and colonization of the Pacific. Cambridge: Cambridge University Press.
10.1017/CBO9780511518225 Google Scholar
- 87 O'Connor S, Ono R, Clarkson C. 2011. Pelagic fishing at 42,000 years before the present and the maritime skills of modern humans. Science 334: 1117–1121.
- 88 O'Connell JF, Allen J, Hawkes K. 2010. Pleistocene Sahul and the origins of seafaring. In: A Anderson et al., editors. The global origins and development of seafaring. Cambridge: McDonald Institute. p 57–68.
- 89 Bailey GN, King GCP, Devès M, et al. 2012. DISPERSE: dynamic landscapes, coastal environments and human dispersals. Antiquity 86: 334. http://antiquity.ac.uk/projgall/bailey334.
- 90 Kusky T, Robinson C, El-Baz F. 2005. Tertiary-Quaternary faulting and uplift in the northern Oman Hajar mountains. J Geol Soc 162: 871–888.
- 91 Bailey GN, Devès M, Inglis RH, et al. n.d. Blue Arabia: Palaeolithic and underwater survey in SW Saudi Arabia and the role of coasts in Pleistocene dispersal. Quat Int. In press. doi:10.1016/j.quaint.2015.01.002.
- 92 Aubert M, Brumm A, Ramli M, et al. 2014. Pleistocene cave art from Sulawesi, Indonesia. Nature 514: 223–227.
- 93 Behar DM, van Oven M, Rosset S, et al. 2012. A “Copernican” reassessment of the human mitochondrial DNA tree from its root. Am J Hum Genet 90: 675–684.
- 94 Gronau I, Hubisz MJ, Gulko B, et al. 2011. Bayesian inference of ancient human demography from individual genome sequences. Nat Genet 43: 1031–1034.
- 95 Hudson RR. 2002. Generating samples under a Wright-Fisher neutral model of genetic variation. Bioinformatics 18: 337–338.
- 96 Fenner JN. 2005. Cross-cultural estimation of the human generation interval for use in genetics-based population divergence studies. Am J Phys Anthropol 128: 415–423.
- 97 Crassard R, Petraglia MD, Drake NA, et al. 2013. Middle Palaeolithic and Neolithic occupations around Mundafan palaeolake, Saudi Arabia: implications for climate change and human dispersals. PLoS ONE 8: e69665.
- 98 Lehner B, Verdin K, Jarvis A. 2008. New global hydrography derived from spaceborne elevation data. EOS, Trans Am Geophys Union 89: 93–94.
10.1029/2008EO100001 Google Scholar
- 99 Breeze P, Drake NA, Groucutt HS, et al. n.d. Remote sensing and GIS techniques for reconstructing Arabian palaeohydrology and identifying archaeological sites. Quat Int. In press. doi:10.1016/j.quaint.2015.01.022.
10.1016/j.quaint.2015.01.022 Google Scholar
- 100 Hijmans RJ, Cameron SE, Parra JL, et al. 2005. Very high resolution interpolated climate surfaces for global land areas. Int J Climatol 25: 1965–1978.
- 101 Jennings RP, Singarayer J, Stone E, et al. n.d. The greening of Arabia: an ensemble of climate model simulations infers multiple opportunities for human occupation of the Arabian Peninsula during the Late Pleistocene. Quat Int. In press. doi:10.1016/j.quaint.2015.01.006.
10.1016/j.quaint.2015.01.006 Google Scholar
- 102 Drake NA, Blench MA, Armitage SJ, et al. 2011. Ancient watercourses and biogeography of the Sahara explain the peopling of the desert. Proc Natl Acad Sci USA 108: 458–462.
- 103 Ambrose SH. 1998. Chronology of the Later Stone Age and food production in East Africa. J Archaeol Sci 25: 377–392.
- 104 Bretzke K, Conard NJ, Uerpmann H-P. 2014. Excavations at Jebel Faya: the FAY-NE1 shelter sequence. Proc Sem Arabian Stud 44: 69–82.
- 105 Nishiaki Y, Kanjo Y, Muhesen S, et al. 2012. The temporal variability of Late Levantine Mousterian lithic assemblages from Dederiyeh Cave, Syria. Eurasian Prehist 9: 3–27.