Genetic and archaeological perspectives on the initial modern human colonization of southern Asia Paul Mellarsa,b,1, Kevin C. Goric,d, Martin Carre,f, Pedro A. Soaresg, and Martin B. Richardse,f,1 aDepartment of Archaeology, University of Cambridge, Cambridge CB2 3DZ, United Kingdom; bDepartment of Archaeology, University of Edinburgh, Edinburgh EH8 9AG, United Kingdom; cDepartment of Biology, University of York, York YO10 5DD, United Kingdom; dEuropean Bioinformatics Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge CB10 1SD, United Kingdom; eArchaeogenetics Research Group, Division of Biology, School of Applied Sciences, University of Huddersfield, Huddersfield HD1 3DH, United Kingdom; fSchool of Biology, Faculty of Biological Sciences, University of Leeds, Leeds LS2 9JT, United Kingdom; and gInstituto de Patologia e Imunologia Molecular da Universidade do Porto, 4200-465 Porto, Portugal Edited by Richard G. Klein, Stanford University, Stanford, CA, and approved May 8, 2013 (received for review April 1, 2013) It has been argued recently that the initial dispersal of anatomically Paleolithic tradition” (1)], broadly similar to those documented modern humans from Africa to southern Asia occurred before the across a wide region of both Europe and western Asia over volcanic “supereruption” of the Mount Toba volcano (Sumatra) at a similar span of time (18). The claim that these industries provide ∼74,000 y before present (B.P.)—possibly as early as 120,000 y B.P. evidence for an early arrival of modern humans from Africa before We show here that this “pre-Toba” dispersal model is in serious 74 ka rests crucially on analyses of two small samples of residual “ ” conflict with both the most recent genetic evidence from both Africa core forms recovered from below the Toba ash-fall deposits (at and Asia and the archaeological evidence from South Asian sites. We Jwalapuram sites 3 and 22), compared with those from a broadly present an alternative model based on a combination of genetic contemporaneous range of Middle Paleolithic/Middle Stone Age analyses and recent archaeological evidence from South Asia and (MSA) sites in other regions of Eurasia and Africa (1, 4, 6). For reasons discussed fully in Archaeology, we regard these analyses as Africa. These data support a coastally oriented dispersal of modern fl ∼ – seriously awed on both archaeological and methodological humans from eastern Africa to southern Asia 60 50 thousand grounds, and the interpretation originally offered for these years ago (ka). This was associated with distinctively African micro- “ ” assemblages has since been withdrawn by the author responsible lithic and backed-segment technologies analogous to the African for the lithic analyses, who “now thinks they might be the work of “ ” GENETICS Howiesons Poort and related technologies, together with a range an unidentified population of archaic people” (ref. 11, p. 26). of distinctively “modern” cultural and symbolic features (highly In archaeological terms, the most remarkable feature of this shaped bone tools, personal ornaments, abstract artistic motifs, model lies in the claims for a rapid and abrupt evolution from microblade technology, etc.), similar to those that accompanied these initial “Indian Middle Paleolithic” industries into the im- the replacement of “archaic” Neanderthal by anatomically modern mediately ensuing “Indian microlithic tradition”, which appears human populations in other regions of western Eurasia at a broadly over effectively all regions of India from at least 35–40 ka onward similar date. [in calibrated radiocarbon terms (19)]—a transformation that, according to the recently dated industrial sequences at Jwala- India | Paleolithic | archaeogenetics | mtDNA puram, occurred within a space of ∼3,000 y, between ∼38 and 35 ka (1–3) (Fig. 1 and Archaeology). According to the evidence ANTHROPOLOGY uestions surrounding the character and chronology of the recorded from the Jwalapuram 9 rock shelter and the two sites of dispersal of anatomically modern human populations from Batadomba-lena and Fahien-lena in Sri Lanka (3, 20, 21) (Fig. 2), Q this allegedly in situ transformation affected virtually all aspects of their African origins ∼150–200 thousand years ago (ka) are cur- the archaeological assemblages, including the sudden appearance rently a focus of great interest and debate. Issues surrounding the of highly controlled microblade-producing techniques (and as- initial modern human colonization of the geographically pivotal sociated core forms), and the production of a wide range of highly region of southern Asia have generated intense controversy in the shaped “microlithic” tools, typical end scrapers, extensively sha- recent genetic and archaeological literature (1–13). fl ped bone tools, a variety of circular, rotary-perforated bead forms, There are currently two sharply con icting models for the ear- and [at the site of Patne in northern India, radiocarbon dated to liest modern human colonization of South Asia, with radically ∼30 ka calibrated (cal.) before present (B.P.)] a fragment of different implications for the interpretation of the associated ge- ostrich eggshell engraved with a complex “bounded criss-cross” netic and archaeological evidence (Fig. 1). The first is that modern – ∼ – design (2, 3, 20 22) (Figs. 3 and 4). At the site of Batadomba- humans arrived 50 60 ka, as part of a generalized Eurasian dis- lena in Sri Lanka these industries are associated with typically persal of anatomically modern humans, which spread (initially as anatomically modern skeletal remains, dated to ∼30–35 ka cal. a very small group) from a region of eastern Africa across the B.P. (21, 23). mouth of the Red Sea and expanded rapidly around the coastlines In all other regions of Europe and western Asia, these ranges of of southern and Southeast Asia, to reach Australia by ∼45–50 ka technological and other cultural innovations are generally seen as (7–10, 14–18) (Fig. 2). The second, more recently proposed view, is the hallmarks of fully “Upper Paleolithic” technologies, which first that there was a much earlier dispersal of modern humans from appeared in the different regions in close association with the ar- Africa sometime before 74 ka (and conceivably as early as 120–130 rival of the first anatomically modern populations from Africa (and ka), reaching southern Asia before the time of the volcanic the associated extinction of the preceding “archaic” Neanderthal “supereruption” of Mount Toba in Sumatra (the largest volcanic populations) ∼45–35 ka cal. B.P. (18, 24–26)—i.e., at a broadly eruption of the past 2 million y) at ∼74 ka (1–6). The arguments advanced in support of the second (“pre-Toba”) colonization model at present hinge on archaeological evidence Author contributions: P.M. and M.B.R. designed research; P.M., K.C.G., M.C., P.A.S., and recovered from recent excavations in the Jwalapuram region of M.B.R. performed research; P.M., K.C.G., M.C., P.A.S., and M.B.R. analyzed data; and P.M. southeastern India (Fig. 2), where a series of stone tool assemblages and M.B.R. wrote the paper. have been recovered from locations both underlying and overlying The authors declare no conflict of interest. thick deposits of Toba ash-fall deposits, with a series of associated This article is a PNAS Direct Submission. ∼ optically stimulated luminescence (OSL) dates ranging from 77 1To whom correspondence may be addressed. E-mail: [email protected] or m.b.richards@ to ∼38 ka (1–6) (Archaeology). Technologically, all of these hud.ac.uk. assemblages are of characteristically Middle Paleolithic form This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. [attributed by the excavators to a generalized “Indian Middle 1073/pnas.1306043110/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1306043110 PNAS Early Edition | 1of6 Downloaded by guest on September 29, 2021 earliest plausible age estimate for the initial modern human col- Petraglia et al. Mellars et al. onization of the region (Fig. 5 and Genetics and Archaeology). ka Archaic humans Modern humans ka i) mtDNA: All extant human populations with deep maternal ancestry in Eurasia carry mtDNA lineages that fall within a sin- 10 10 gle haplogroup, L3, which most likely originated initially in eastern Africa (8, 14, 27, 28). L3 diverged during the dispersal 20 20 from Africa into Asia into the derived M, N, and R lineages Microlithic (10, 15, 29, 30). Because the makers of the microlithic technol- Microlithic industries industries (Modern humans) ogies of India are agreed to provide the demographic basis for 30 30 most if not all of the present-day Indian populations (see above), they almost certainly carried the M and N/R lineages Microlithic ? 40 40 transition that today account for all known South Asian mtDNA lineages (excepting recent immigrants). In the context of the present ? Coastal sites discussion the critical issue is whether these lineages could 50 50 Modern humans Middle plausibly have arrived in India before the Toba eruption Palaeolithic (∼74 ka). industries 60 60 Middle Palaeolithic The lineages have been dated in a number of ways, with (Archaic humans) varying assumptions, but the earliest plausible estimates cur- 70 70 TOBA rently fall at ∼70 ka (95% range: 61.6–79 ka) for L3 in Africa ERUPTION (the upper bound for departure), 61 ka (50.4–72.1 ka) for the ? Indian AMH 80 80 Colonisation most ancient non-African haplogroup, haplogroup N, in Arabia, and 48 ka (39.6–56.5 ka) and 62.3 ka (54.7–70 ka) for Fig. 1. Comparison of the two alternative models for the initial modern M and R in South Asia [Fig. 5 and Table S1 (8)]. (Note that R human colonization of South Asia discussed in the text. The graphs show the must be slightly younger than N, because R nests within N, inferred correlations between “archaic” and “modern” populations and which is allowed for given the errors on these estimates.) An their associations with Middle Paleolithic vs.