1. Going East: New Genetic and Archaeological
Perspectives on the Modern Human Colonization of
Eurasia
Paul Mellars, et al.
Science 313, 796 (2006);
DOI: 10.1126/science.1128402
The following resources related to this article are available online at
www.sciencemag.org (this information is current as of July 19, 2007 ):
Updated information and services, including high-resolution figures, can be found in the online
version of this article at:
http://www.sciencemag.org/cgi/content/full/313/5788/796
A list of selected additional articles on the Science Web sites related to this article can be
Downloaded from www.sciencemag.org on July 19, 2007
found at:
http://www.sciencemag.org/cgi/content/full/313/5788/796#related-content
This article cites 40 articles, 6 of which can be accessed for free:
http://www.sciencemag.org/cgi/content/full/313/5788/796#otherarticles
This article has been cited by 5 article(s) on the ISI Web of Science.
This article has been cited by 2 articles hosted by HighWire Press; see:
http://www.sciencemag.org/cgi/content/full/313/5788/796#otherarticles
This article appears in the following subject collections:
Anthropology
http://www.sciencemag.org/cgi/collection/anthro
Information about obtaining reprints of this article or about obtaining permission to reproduce
this article in whole or in part can be found at:
http://www.sciencemag.org/about/permissions.dtl
Science (print ISSN 0036-8075; online ISSN 1095-9203) is published weekly, except the last week in December, by the
American Association for the Advancement of Science, 1200 New York Avenue NW, Washington, DC 20005. Copyright
c 2006 by the American Association for the Advancement of Science; all rights reserved. The title SCIENCE is a
registered trademark of AAAS.

2. MIGRATION AND DISPERSAL
result that could allow us to link ultimate and 3. J. R. Busvine, Disease Transmission by Insects: Its 25. E. G. Oliveira, R. B. Srygley, R. Dudley, J. Exp. Biol. 201,
proximate aspects of insect migration (14, 36). Discovery After 90 Years of Effort to Prevent It (Springer- 3317 (1998).
Verlag, Berlin, 1993). 26. H. Mouritsen, B. J. Frost, Proc. Natl. Acad. Sci. U.S.A. 99,
Such studies may also provide new insights into 4. C. Kremen, R. S. Ostfeld, Front. Ecol. Environ. 3, 540 (2005). 10162 (2002).
the genetic control of migratory flight decisions 5. J. Roffey, J. I. Magor, ‘‘Desert locust population param- 27. R. B. Srygley, R. Dudley, E. G. Oliveira, A. J. Riveros,
per se (9). Biomechanical models can be used to eters’’ (Food and Agriculture Organization of the United Anim. Behav. 71, 183 (2006).
improve our understanding of the energetic costs Nations, Rome, 2003). 28. R. B. Srygley, E. G. Oliveira, R. Dudley, Proc. R. Soc.
6. P. Berthold, Bird Migration: A General Survey (Oxford London B. Biol. Sci. 263, 1351 (1996).
of insect migration (25, 27–29), and the trade-offs Univ. Press, Oxford, ed. 2, 2001). 29. R. B. Srygley, E. G. Oliveira, J. Navig. 54, 405 (2001).
those costs impose on other aspects of an insect’s 7. H. Dingle, Migration: The Biology of Life on the Move 30. F. A. Urquhart, Proc. Entomol. Soc. Ont. 95, 22
life history (37). Most important, our ability to (Oxford Univ. Press, Oxford, 1996). (1965).
track small animals over large distances is 8. C. B. Williams, Annu. Rev. Entomol. 2, 163 (1957). 31. W. W. Cochran, H. Mouritsen, M. Wikelski, Science 304,
9. L. P. Brower, J. Exp. Biol. 199, 93 (1996). 405 (2004).
improving steadily. Current systems for tracking 10. J. S. Kennedy, in Migration: Mechanisms and Adaptive 32. D. F. Naef-Daenzer, M. Stalder, P. Wetli, E. Weise, J. Exp.
animals globally are unsuitable for creatures Significance, M. A. Rankin, Ed. [Contrib. Mar. Sci. 27 Biol. 208, 4063 (2005).
smaller than È500 g, which excludes the majority (suppl.), 5 (1985)]. 33. P. D. Lorch, G. A. Sword, D. T. Gwynne, G. L. Anderson,
of birds and bats as well as all insects. However, 11. I. P. Woiwod, D. R. Reynolds, C. D. Thomas, Insect Ecol. Entomol. 30, 548 (2005).
Movement: Mechanisms and Consequences (CABI, 34. M. Wikelski et al., Biol. Lett., in press.
the signals from transmitters now being used to
Wallingford, UK, 2001). 35. R. B. Gasser, Vet. Parasitol. 136, 69 (2006).
track dragonflies could be received from space 12. P. C. Gregg, G. P. Fitt, M. P. Zalucki, D. A. H. and Murray, in 36. A. G. Gatehouse, Annu. Rev. Entomol. 42, 475 (1997).
with the installation of a small-animal tracking Insect Migration: Tracking Resources Through Space and 37. A. J. Zera, Integr. Comp. Biol. 45, 511 (2005).
Downloaded from www.sciencemag.org on July 19, 2007
satellite. Such a system is technologically feasible Time, V. A. Drake, A. G. Gatehouse, Eds. (Cambridge Univ. 38. G. W. Swenson, M. Wikelski, J. A. Smith, paper presented
(38). The ability to follow individual insects Press, Cambridge, 1995), pp. 151–172. at the Proceedings of the IEEE International Geoscience
13. T. R. E. Southwood, J. Anim. Ecol. 46, 337 (1977). Remote Sensing Symposium, Anchorage, Alaska, 20 to 24
throughout their migrations will be invaluable to 14. K. Wilson, in Insect Migration: Tracking Resources September 2004.
understanding the selective forces behind insect Through Space and Time, V. A. Drake, A. G. Gatehouse, 39. L. P. Brower, Anim. Kingdom 91, 42 (1988).
migration. We also need data on individual insects Eds. (Cambridge Univ. Press, Cambridge, 1995), 40. H. Aridjis, L. P. Brower, ‘‘Twilight of the monarchs,’’
to understand how migratory abundance changes pp. 243–264. New York Times, 26 January 1996, p. A27.
15. R. W. Russell, M. L. May, K. L. Soltesz, J. W. Fitzpatrick, 41. A. R. E. Sinclair, in Serengeti II: Dynamics, Management,
with climatic cycles and with the use of pesticides Am. Midl. Nat. 140, 325 (1998). and Conservation of an Ecosystem, A. R. E. Sinclair,
on targeted and nontargeted species. Until the 16. V. A. Drake, R. A. Farrow, Annu. Rev. Entomol. 33, 183 (1988). P. Arcese, Eds. (Univ. of Chicago Press, Chicago, 1995).
migratory behavior of individuals can be separated 17. A. G. Gatehouse, X. X. Zhang, in Insect Migration: 42. M. J. Harvey, J. S. Atlenbach, T. L. Best, Bats of the United
out from the behavior of populations, insect Tracking Resources Through Space and Time, V. A. Drake, States (Arkansas Game and Fish Commission, Little Rock,
A. G. Gatehouse, Eds. (Cambridge Univ. Press, AR, 1999).
migration is likely to remain a poorly understood Cambridge, 1995), vol. 193–242. 43. P. A. Johnsgard, Crane Music: A Natural History of
but immensely important phenomenon. 18. R. A. Farrow, in The Biology of the Grasshoppers, American Cranes (Smithsonian Institution, Washington,
R. F. Chapman, A. Joern, Eds. (Wiley, London, 1990), DC, 1991).
References and Notes pp. 227–314. 44. We thank W. W. Cochran for inspiration and the National
1. V. A. Drake, A. G. Gatehouse, Eds., Insect Migration: 19. M. P. J. Shaw, Ann. Appl. Biol. 65, 197 (1970). Geographic Society and the National Science Foundation
Tracking Resources Through Space and Time (Cambridge 20. M. Collett, T. S. Collett, Curr. Biol. 16, R48 (2006). (SGER-0528881) for support. R.A.H is supported by a
Univ. Press, Cambridge, 1995). 21. A. C. Perdeck, Ardea 46, 1 (1958). Marie Curie Outgoing international fellowship (MOIF-CT-
2. J. I. Magor, in Insect Migration: Tracking Resources Through 22. V. P. Bingman, K. Cheng, Ethol. Ecol. Evol. 17, 295 (2006). 2005-021508).
Space and Time, V. A. Drake, A. G. Gatehouse, Eds. 23. L. R. Taylor, J. Anim. Ecol. 43, 225 (1974).
(Cambridge Univ. Press, Cambridge, 1995), pp. 399–426. 24. W. H. Calvert, J. Lepid. Soc. 55, 162 (2001). 10.1126/science.1127272
pecially those based on the maternally inherited
REVIEW mitochondrial DNA) strongly suggest that a
small subset of these African populations made
Going East: New Genetic and the crossing from northeastern Africa, probably
over the mouth of the Red Sea, and subse-
quently dispersed into Arabia and southern Asia
Archaeological Perspectives on the sometime before 50,000 years before present
(yr B.P.) (2, 8, 12–17) (Fig. 1). Recent studies
Modern Human Colonization of Eurasia have suggested that these populations expanded
rapidly along the coastlines of southern Asia,
Paul Mellars southeastern Asia, and Indonesia to arrive in
both Malaysia and the Andaman Islands by at
The pattern of dispersal of biologically and behaviorally modern human populations from their least 55,000 yr B.P., and conceivably as early as
African origins to the rest of the occupied world between È60,000 and 40,000 years ago is at present 60,000 to 65,000 yr B.P. (12, 18–21)—though
a topic of lively debate, centering principally on the issue of single versus multiple dispersals. Here I more recent estimates of mitochondrial DNA mu-
argue that the archaeological and genetic evidence points to a single successful dispersal event, which tation rates (8) suggest that these figures may be
took genetically and culturally modern populations fairly rapidly across southern and southeastern overestimates. As Carl Sauer pointed out in 1962
Asia into Australasia, and with only a secondary and later dispersal into Europe. (22), a strongly coastal pattern of dispersal would
make good sense in ecological and demographic
esearch over the past 20 years has dicate that human populations that were essen- terms, because this would presumably have re-
R provided an increasingly clear picture
of the way in which our own species
(Homo sapiens) emerged and subsequently
tially Bmodern[ both anatomically and in their
mitochondrial and Y-chromosome lineages had
emerged in Africa by at least 150,000 years ago,
quired only limited economic adaptations from
one coastal location to another.
spread across the rest of the occupied world. perhaps closer to 200,000 years ago (1–11). Department of Archaeology, Cambridge University, Cam-
DNA evidence and fossil skeletal remains in- Studies of present-day world populations (es- bridge CB2 3DZ, UK. E-mail: pam59@cam.ac.uk
796 11 AUGUST 2006 VOL 313 SCIENCE www.sciencemag.org

3. SPECIALSECTION
The subsequent dispersal southward of these but possibly two or more separate dispersals modern Asian and European populations derive
populations into New Guinea and Australia (at of anatomically and genetically modern hu- from one small subset of the so-called L3 mito-
that time connected as an extended landmass mans from Africa into Eurasia. An earlier mod- chondrial lineage in Africa, which subsequently
known as BSahul[) is currently more controver- el advanced by Lahr and Foley in 1994 (29–31) diverged into the derivative M and N lineages,
sial. We know that typically anatomically mod- envisaged at least two separate episodes of probably shortly after their dispersal from
ern populations were present at the Niah cave in dispersal from northeastern Africa, associated Africa (2, 8, 12–15). The crux of the arguments
Sarawak by at least 41,000 years ago (23), and with sharply differing patterns of stone-tool advanced by Kivisild (8), Forster (2), Matsumura
that some of these populations had made a sea technology. Of these, the ‘‘northern’’ dispersal (12), Macaulay et al. (18), and others is that
crossing of at least 90 km to reach parts of extended northward via the Nile Valley and the the very limited genetic diversity exhibited
Australia by at least 45,000 yr B.P.—best rep- Sinai Peninsula into southwestern Asia (and even- by modern European and Asian populations—
resented by the typically anatomically modern tually Europe), associated with typically blade- compared to those in Africa—would be effec-
skeleton from the site of Lake Mungo 3 in New dominated, ‘‘Upper Palaeolithic’’ or ‘‘Mode 4’’ tively impossible to reconcile with the model of
South Wales (1, 24–26). Some archaeological technology, best represented at the sites of two separate dispersal events, deriving from sep-
claims have been made for the initial colonization Boker Tachtit in southern Israel and Ksar Akil arate source populations in Africa, and hypo-
of Australia as early 50,000 to 60,000 yr B.P. in Lebanon, both dated to around 45,000 to thetically at two different dates. Essentially the
(27), but these remain speculative and contested 50,000 yr B.P. (32–34). The separate ‘‘southern’’ same conclusion has been drawn independently
(24). Similarly, claims for the presence of behav- dispersal extended from the Horn of Africa by Endicott and others (13) based on studies of
Downloaded from www.sciencemag.org on July 19, 2007
iorally modern populations in Malaysia before across the mouth of the Red Sea (the Bab el the paternally inherited Y-chromosome lineages.
the Mount Toba (Sumatra) volcanic Bsuper- Mandeb straits) carrying technologically simpler Even allowing for the current controversies sur-
eruption[ around 74,000 yr B.P. (16) have yet ‘‘Middle Palaeolithic’’ or ‘‘Mode 3’’ technology, rounding the interpretation and dating of the
to receive any clear support from recent archaeo- which subsequently dispersed eastward along the DNA evidence (2, 5, 8, 11, 14, 35–37), it is be-
logical research in the region (28). But in any coasts of southern and southeastern Asia into coming increasingly difficult to reconcile the
event, it is clear that the initial dispersal of Australia (21, 29–31). The sharp contrasts in the available genetic data with the hypothesis of two
modern human populations eastward from technology associated with these two dispersals or more separate dispersal events from Africa
their original African homeland along the were taken as an explicit reflection of two sep- into Eurasia—although this point has been de-
so-called coastal express (12, 18, 20) route into arate source populations in Africa, with the bated in some earlier genetic studies (15, 38).
Australasia occurred over a comparatively short southern, Mode 3 dispersal occurring substan-
time, amounting to at most 15,000 years, and tially earlier than the northern, Mode 4 dis- The Archaeological Evidence
probably less than 10,000 years (i.e., an overall persal event (29–31). The major challenge of this scenario now is to
dispersal rate of at least 1.0 km per year) if we Recently, the notion of two or more separate document the individual steps in this coloniza-
take the combined DNA and archaeological dispersals of anatomically and genetically mod- tion process on the basis of the ‘‘hard’’ archaeo-
evidence into account (12, 18, 21). ern humans has come under increasing scrutiny logical evidence. Large areas of both Arabia and
from molecular geneticists, based on studies of India in particular are at present largely blank
Single or Multiple Dispersals? both mitochondrial and Y-chromosome patterns areas on the archaeological map over the crit-
The most controversial issue at present cen- in African and Asian populations (2, 8, 12–15, 18). ical time range from È50,000 to 60,000 yr B.P.
ters on whether there could have been not one, These recent studies suggest that the whole of in question (39, 40). And of course, all the coast-
lines of this period are now deeply submerged
below the rapidly rising sea levels of the past
15,000 years (20, 21). There are, however, al-
ready some intriguing hints from south Asia of
what future research may reveal. From the sites
of Patne in western India (41), Jwalapuram in
southeast India (42), and Batadomba-lena in Sri
Lanka (43, 44), there are archaeological assem-
blages showing some striking resemblances to
those from eastern and southern Africa that must
be from very close to the period when modern
humans first dispersed from Africa (Fig. 2A).
These sites contain large numbers of small
‘‘crescentic’’ forms of stone tools (evidently parts
of hafted implements, and conceivably compo-
nents of archery equipment) (45, 46) that are
markedly similar to those that define the so-
called Howiesons Poort technology in southern
and eastern Africa, dated broadly to around
55,000 to 65,000 yr B.P. (45–51)—as at the sites
of Mumba in Tanzania (45, 52), Norikiushin in
Kenya (53), and a range of similar sites in
southern Africa (45–49) (Fig. 2B). Broadly
Fig. 1. Map of possible dispersal routes of anatomically and genetically modern human pop- similar industries including rather larger forms
ulations from Africa to Asia and Australia according to Forster and Matsumura (12). The models of backed ‘‘segment’’ forms are dated to be-
assume an origin in eastern Africa, and dispersal either via the Nile Valley and Sinai Peninsula (the tween 60,000 and 40,000 yr B.P. at the site of
‘‘northern’’ route) or via the mouth of the Red Sea to Arabia and Australia (the ‘‘southern’’ route). Enkapune ya Muto in Kenya (53, 54). Although
www.sciencemag.org SCIENCE VOL 313 11 AUGUST 2006 797

4. MIGRATION AND DISPERSAL
the Indian crescentic tools are generally smaller these striking similarities in material culture are as end scrapers, backed blades, or burins—a
than those from the African sites, the sequences entirely coincidental, they point strongly to a di- pattern of technology that persisted in Australia
at Patne and Jwalapuram suggest that these rect connection between the earliest modern hu- from at least 45,000 yr B.P. down to the middle
forms become increasingly larger in the lower man colonists in southern Asia and their probable of the Holocene period, around 5000 to 7000 yr
levels of the sequence, and correspondingly more ancestors in eastern and southern Africa. B.P. (26, 57, 58). How can we reconcile this
similar to the African tools (41, 42). Even more observation with the hypothesis that these
strikingly, these Indian sites have produced The Australian Archaeological Record technologies developed from more ‘‘advanced,’’
carefully shaped and perforated beads manu- The greatest enigma in the current archae- blade-based technologies in Africa (and appar-
factured from fragments of ostrich eggshell, ological record lies in the lack of similarly ently parts of India) with the initial dispersal of
closely similar to those found in African sites ‘‘advanced’’ technologies in the areas to the east anatomically and genetically modern populations
(39, 41, 45, 47), together with a further piece of of the Indian subcontinent, and especially in the in their eastward migration?
ostrich eggshell incised with a distinctive criss- relatively well-explored areas of Australia and The answer to this paradox might lie partly
cross motif (41), which is strongly reminiscent New Guinea, which were colonized by anatom- in environmental factors, and partly in the pat-
of designs engraved on pieces of red ochre from ically modern humans from at least 45,000 yr terns of cultural and technological development
the later Middle Stone Age levels in the B.P. onward, as discussed above (24–26). The that are probably inherent in the progressive
Blombos Cave in South Africa (55) (Fig. 2, A earliest stone-tool technologies documented dispersal of small-scale human populations
and B), together with similar designs incised on across the whole of Australasia are conspicu- across a long and environmentally complex colo-
Downloaded from www.sciencemag.org on July 19, 2007
fragments of ostrich-eggshell water containers ously lacking in any trace of distinctively nization route. Three factors in particular are
from the site of Diepkloof in the western Cape ‘‘modern’’ or ‘‘Upper Palaeolithic,’’ blade-based likely to have been significant in this context.
(56), dated respectively to È75,000 and 60,000 technologies of the kind recorded from both The first, and potentially most important, factor
yr B.P. (50, 51). At present, the Indian and Sri the later African Middle Stone Age sites and lies in the general scarcity of high-quality, fine-
Lankan sites in question can only be reliably the earliest modern human sites in southwest grained stone for tool production in most areas of
dated back to around 34,000 yr B.P. (in cal- Asia and Europe (26, 57, 58). These Australian eastern and southeastern Asia (59, 60). Blade
ibrated radiocarbon terms) (34, 39, 41–44), but technologies consist of very simple, flake-based technology in particular is heavily dependent
current excavations at the Jwalapuram site in industries, completely lacking in typical blade on the availability of nodules of fine-grained
southern India suggest that similar technologies forms and apparently with little or no trace of stone such as flint, chert, obsidian, or other
may go back to a much earlier date (42). Unless typically Upper Palaeolithic tool forms such fine-grained rocks, which are scarce over many
Fig. 2. (A) (Upper) Assemblages of ‘‘crescentic’’ and related stone tool respectively. (B) (Upper) Crescentic and related tool forms from the
forms from the sites of Patne in western India (41) and Batadomba-lena in ‘‘Howieson’s Poort’’ levels at Klasies River (South Africa) dated to È60,000
Sri Lanka (43), radiocarbon dated to between 30,000 and 34,000 to 65,000 yr B.P. (48). (Lower) Geometric designs incised on pieces of red
(calibrated) yr B.P. (Lower) Geometric design engraved on ostrich eggshell ochre from the Blombos Cave, South Africa (È75,000 yr B.P.) (55) and
fragment from Patne (left) and specimens of ostrich eggshell beads (and ostrich eggshell beads from the site of Enkapune ya Muto, Kenya (È40,000
preforms) and a perforated shell recovered from Patne and Batadomba-lena, yr B.P.) (56).
798 11 AUGUST 2006 VOL 313 SCIENCE www.sciencemag.org

5. SPECIALSECTION
areas of eastern Asia. Even if there were oc- complexity and diversity of cultural and tech- Australian industries (such as those from Lake
casionally localized areas of better-quality nological patterns with increasing distance from Mungo in New South Wales) (26, 57, 58, 68)
stone, such as obsidian or high-quality cherts, their demographic point of origin. Arguably the have more in common with simplified forms of
the patterns of lithic technology would inevita- most striking illustration of this kind of founder single-platform blade cores (Fig. 4) in their
bly have to ‘‘adapt’’ to the more extensive areas effect and technological drift process can be basic conceptual and flaking strategies than
in which these better-quality raw materials were seen in the loss of several technological fea- with the distinctively ‘‘radial’’ or ‘‘centripetal’’
lacking. Indeed, it has often been suggested that tures (such as fishing, bone tools, and other patterns of flaking, which are the hallmark of
the technology over large areas of eastern Asia cultural elements) associated with the settle- the African and Eurasian Middle Palaeolithic
may have been much more dependent on wood, ment and cultural development of Tasmanian prepared core techniques (59, 69). Any sugges-
or even bamboo, for tool manufacture than on populations, following their initial colonization tion that the earliest colonists of southeast Asia
conventional stone tool supplies (59, 60). of the island around 35,000 yr B.P. (64–66). and Australia carried with them distinctively
The second major factor in stone tool All of these cultural and technological pro- Middle Palaeolithic, Mode 3 technologies, as
technology lies in the specific functions for cesses could be seen as a direct parallel to the suggested in the ‘‘multiple dispersals’’ model
which the tools were required. If, as most of the progressive loss in the genetic diversity of (29–31, 39), would seem to have little support
current models suggest, the initial colonization the dispersing modern human populations in the documented archaeological record from
of southeastern Asia and Australasia followed a over geographical trajectories extending from Australia. The totally ‘‘modern’’ character of
primarily coastal route (12, 18, 20, 21, 61), then their putative African origin progressively east- the burial rituals, personal ornaments, abundant
Downloaded from www.sciencemag.org on July 19, 2007
the technologies would be likely to adapt pri- ward and westward—as recently documented use of red ochre, and elaborate ground and
marily to the exploitation of coastal resources, in studies of present-day genetic patterns by shaped stone axes, documented from effectively
such as fish, shellfish, and marine mammals Prugnolle et al. (6), Liu et al. (7), and others the earliest stages of colonization of Austra-
(together with tropical plant foods) with per- (Fig. 3). lia (26, 57, 58, 70), should also be kept in
haps only a minor component of hunting larger When viewed in these terms, the relative mind in this context. As several authors have
land mammals, of the kind that clearly formed ‘‘simplicity’’ of the technology associated with pointed out in relation to the 100,000-year-old
a major part of the human economy in both the initial modern human settlement of south- ritualistic burials from the sites of Skhul and
Africa and the whole of western Asia and Europe east Asia and Australia becomes not merely Qafzeh in Israel (17, 71–73), there is clearly
(21, 59). This would presumably have involved plausible but arguably largely predictable, in much more to the emergence of cognitively
much less emphasis on various forms of hunting demographic and cultural terms. Although these ‘‘modern,’’ symbolically constructed behavior
equipment (such as spears, meat-processing early Australian technologies are strikingly than the production of typically Upper Palaeo-
tools, etc.), as well as equipment involved in the different from those of the early Upper Palaeo- lithic stone tools.
manufacture of elaborate skin clothing, or the lithic, blade-dominated industries in south-
construction of tents and other living structures, western Asia and Europe, there is nothing The Colonization of Western Asia and Europe
that were essential to survival in much colder, distinctively Middle Palaeolithic (or Mode 3) One important implication of this single-dispersal,
more northerly environments (59, 62). about their character (29–31). The whole of the ‘‘southern route’’ colonization of Eurasia from
Finally, there is the more fundamental Australian technology is conspicuously lacking
demographic and evolutionary issue of the in anything resembling typically Middle Palaeo-
repeated founder effects, and associated cultural lithic ‘‘Levallois’’ or similar ‘‘prepared core’’ tech-
drift, as relatively small population units ex- niques (26, 57, 58), and the kinds of simple
panded progressively eastward along the south- retouched flake tools encountered in these
ern and southeastern Asian coasts (21, 63). These industries could be paralleled just as easily in
repeated, successive, and cumulative small-scale some of the early Upper Palaeolithic industries
founder effects would inevitably operate not in Europe (67) as in the Eurasian and African
only on the biological and genetic features of the Middle Palaeolithic/Middle Stone Age sites.
populations (6, 7, 11) but also on their reper- More specifically, it could be argued that the
toires of cultural and technological behavior— curious, single-platform ‘‘horse-hoof’’ cores
probably leading to a progressive loss in the that are such a distinctive feature of the earliest
Fig. 4. ‘‘Horse hoof’’ core from Lake Mungo,
Australia, dated to È40,000 to 45,000 B.P. The
Fig. 3. Decline in the genetic diversity of present-day human populations with increasing distance basic flaking strategy of these cores is analogous to
from the presumed point of dispersal of anatomically and genetically modern populations in East that of simplified forms of single-platform blade
Africa. [Reprinted from Prugnolle et al. (6), with permission from Elsevier.] cores (26).
www.sciencemag.org SCIENCE VOL 313 11 AUGUST 2006 799

6. MIGRATION AND DISPERSAL
eastern Africa is that the modern human evidence, the finer details of this colonization 44. A. R. Kennedy et al., Am. J. Phys. Anthropol. 72, 441
process have still to be worked out. (1987).
colonization of the Near East and Europe must
45. S. McBrearty, A. Brooks, J. Hum. Evol. 39, 453 (2000).
have reached these areas via western or southern 46. P. Mellars, in The Speciation of Modern Homo Sapiens,
Asia, rather than via the Nile Valley and Sinai References and Notes T. J. Crow, Ed. (British Academy, London, 2002), pp. 31–47.
Peninsula, as implied in the ‘‘northern’’ dispersal 1. C. B. Stringer, Philos. Trans. R. Soc. London Ser. B 357, 47. H. J. Deacon, J. Deacon, Human Beginnings in South
563 (2002). Africa: Uncovering the Secrets of the Stone Age (David
route. This conclusion receives further support 2. P. Forster, Philos. Trans. R. Soc. London Ser. B 359, 255 Philip, Cape Town, South Africa, 1999).
from the relatively late dating of the arrival of (2004). 48. R. Singer, J. Wymer, The Middle Stone Age at Klasies River
modern humans in Europe (È40,000 to 45,000 3. T. D. White et al., Nature 423, 742 (2003). Mouth in South Africa (Univ. of Chicago Press, Chicago,
yr B.P.) implied by both the archaeological and 4. I. McDougall et al., Nature 433, 733 (2005). 1982).
5. S. A. Tishkoff, S. M. Williams, Nat. Rev. Genet. 3, 611 49. S. Wurz, S. Afr. Archaeol. Bull. 54, 38 (1999).
DNA evidence (9, 34, 74, 75), as compared to (2002). 50. J. K. Feathers, J. Archaeol. Sci. 29, 177 (2002).
the earlier colonization of southern and south- 6. F. Prugnolle, A. Manica, F. Balloux, Curr. Biol. 15, R159 51. C. Tribolo, N. Mercier, H. Valladas, in From Tools to
eastern Asia implied by both the DNA evidence (2005). Symbols: From Early Hominids to Modern Humans,
and the early archaeological and skeletal 7. H. Liu, F. Prugnolle, A. Manica, F. Balloux, Am. J. Hum. F. d’Errico, L. Blackwell, Eds. (Wits Univ. Press,
Genet. 79, 230 (2006). Johannesburg, South Africa, 2005), pp. 493–511.
evidence from Australia. Some archaeological 52. M. J. Mehlman, in Cultural Beginnings: Approaches to
8. T. Kivisild et al., Genetics 172, 373 (2006).
support for the northern route could perhaps be 9. P. Underhill et al., Ann. Hum. Genet. 65, 43 (2001). Understanding Early Hominid Lifeways in the African
seen in the presence of early blade industries at 10. L. Ray, M. Currat, P. Berthier, L. Excoffier, Genome Res. Savannah, J. D. Clark, Ed. (Habelt, Bonn, 1991),
sites such as Taramsa in the Nile Valley (76), 15, 1161 (2006). pp. 177–196.
Downloaded from www.sciencemag.org on July 19, 2007
11. M. A. Jobling, M. Hurles, C. Tyler-Smith, Human 53. S. H. Ambrose, in Thinking Small: Global Perspectives on
but these sites are only tentatively dated at
Evolutionary Genetics: Origins, Peoples and Disease Microlithic Technologies, R. Elston, S. L. Kuhn, Eds.
present, and the industries show at best only (Garland Science, New York, 2004). (American Anthropological Association, Washington, DC,
tenuous morphological similarities to those 12. P. Forster, S. Matsumura, Science 308, 965 (2005). 2004), pp. 9–29.
from the 45,000- to 50,000-year-old sites of 13. P. Endicott, M. Metspalu, T. Kivisild, in South Asia at the 54. S. H. Ambrose, J. Archaeol. Sci. 25, 377 (1998).
Boker Tachtit and Ksar Akil in the Near East Crossroads, M. Petraglia, B. Allchin, Eds. (Springer, 55. C. S. Henshilwood et al., Science 295, 1278 (2002).
Heidelberg, Germany, in press). 56. J. Parkington, C. Poggenpoel, J.-P. Rigaud, P. Texier, in
(32, 34, 76). 14. M. Metspalu et al., BMC Genet. 5, 26 (2004). From Tools to Symbols: From Early Hominids to Modern
The alternative possibility would be to see 15. L. Quintana-Murci et al., Am. J. Hum. Genet. 74, 827 Humans, F. d’Errico, L. Blackwell, Eds. (Wits Univ. Press,
the origins of the Near Eastern industries as (2004). Johannesburg, South Africa, 2005), pp. 475–492.
related directly to the single, southern dispersal 16. S. Oppenheimer, The Peopling of the World (Constable, 57. J. Flood, Archaeology of the Dreamtime: The Story of
London, 2003). Aboriginal Australia and Its People (Collins, London, 1983).
event, which carried modern humans first 17. P. Mellars, Proc. Natl. Acad. Sci. U.S.A. 103, 9381 58. H. Lourandos, Continent of Hunter-Gatherers:
across the Bab el Mandeb straits to Arabia, (2006). New Perspectives in Australian Prehistory (Cambridge
followed by a splitting of the migration routes, 18. V. Macaulay et al., Science 308, 1034 (2005). Univ. Press, Cambridge, 1997).
one moving eastward toward India, and the 19. K. Thangaraj et al., Science 308, 996 (2005). 59. R. G. Klein, The Human Career (Chicago Univ. Press,
20. C. Stringer, Nature 405, 24 (2000). Chicago, 1999).
other moving northward, either through central 21. J. S. Field, M. M. Lahr, J. World Prehist. 19, 1 (2006). 60. K. D. Schick, N. Toth, Making Silent Stones Speak:
Arabia or the eastern coast of the Red Sea into 22. C. Sauer, Proc. Am. Philos. Soc. 106, 41 (1962). Human Evolution and the Dawn of Technology
the Near East (8, 13, 14, 16, 21)—with an as- 23. G. Barker et al., Proc. Prehist. Soc. 68, 147 (2002). (Phoenix, London, 1995).
sociated divergence of technological patterns 24. J. F. O’Connell, J. Allen, J. Archaeol. Sci. 31, 835 61. S. Bowdler, in Sunda and Sahul, J. Allen, G. Golson, R. Jones,
(2004). Eds. (Academic Press, London, 1977), pp. 204–246.
during the course of these dispersals. Alterna-
25. J. M. Bowler et al., Nature 421, 837 (2003). 62. C. W. Marean, in From Tools to Symbols: From Early
tively, this split could have occurred rather 26. J. Mulvaney, J. Kamminga, Prehistory of Australia Hominids to Modern Humans, F. d’Errico, L. Blackwell, Eds.
further to the east in, say, eastern Arabia or Iran (Allen & Unwin, St. Leonards, NSW, Australia, 1999). (Wits Univ. Press, Johannesburg, South Africa, 2005), pp.
(14, 16, 21). The character of the industry from 27. R. Jones, Proc. Br. Acad. 99, 37 (1998). 333–371.
the basal (‘‘Endingi’’) levels in the Enkapune 28. F. Bulbeck, in Under the Canopy: The Archaeology of 63. V. Eswaran, Curr. Anthropol. 43, 749 (2002).
Tropical Rain Forests, J. Mercader, Ed. (Rutgers Univ. 64. R. Jones, in Explorations in Ethnoarchaeology, R. Gould, Ed.
ya Muto rock shelter in Kenya, dated to Press, NJ, 2003). (Univ. of New Mexico, Albuquerque, NM, 1977), pp. 11–47.
È50,000 to 60,000 yr B.P. (comprising elon- 29. M. M. Lahr, R. Foley, Evol. Anthropol. 3, 48 (1994). 65. J. Diamond, Nature 273, 185 (1978).
gated Levallois points and blade forms, remi- 30. R. Foley, M. M. Lahr, Camb. Archaeol. J. 7, 3 (1997). 66. J. Henrich, Am. Antiq. 69, 197 (2004).
niscent of those in the earliest Near Eastern 31. M. M. Lahr, R. Foley, Yearb. Phys. Anthropol. 41, 127 67. D. de Sonneville-Bordes, Le Paleolithique Superieur en
´ ´
(1998). Perigord (Delmas, Bordeaux, France, 1960).
´
industries) (54), could perhaps provide some 32. O. Bar-Yosef, in The Geography of Neandertals and 68. W. Shawcross, Archaeol. Oceania 33, 103 (1998).
support for this model. Modern Humans in Europe and the Greater Mediterranean, 69. P. Mellars, The Neanderthal Legacy: An Archaeological
But until the character of the archaeological O. Bar-Yosef, D. Pilbeam, Eds. (Peabody Museum, Perspective from Western Europe (Princeton Univ. Press,
evidence from both the Nile Valley and Arabia Cambridge, MA, 2000). Princeton, NJ, 1996).
33. P. Mellars, Nature 432, 461 (2004). 70. A. Brumm, M. W. Moore, Camb. Archaeol. J. 15, 157
is better documented (40, 76), the precise source 34. P. Mellars, Nature 439, 931 (2006). (2005).
of the earliest modern human colonists in the 35. V. Eswaran, H. Harpending, A. R. Rogers, J. Hum. Evol. 71. E. Hovers, S. Ilani, O. Bar-Yosef, B. Vandermeersch,
Near Eastern zone will be difficult to pin down. 49, 1 (2005). Curr. Anthropol. 44, 491 (2003).
And the evidence for the earlier, unsuccessful 36. H. Harpending, V. Eswaran, Science 309, 1995 72. J. J. Shea, Evol. Anthropol. 12, 173 (2003).
(2005). 73. M. Vanhaeren et al., Science 312, 1786 (2006).
dispersal of anatomically modern humans from
37. D. B. Goldstein, L. Chikhi, Annu. Rev. Genom. Hum. 74. P. Mellars, Evol. Anthropol., in press.
North Africa into the Near East around 100,000 Genet. 3, 129 (2002). 75. M. Richards et al., Am. J. Hum. Genet. 67, 1251 (2000).
years ago (as represented by the finds from 38. L. Quintana-Murci et al., Nat. Genet. 23, 437 (1999). 76. P. Van Peer, Anthropologie (Brno) 42, 241 (2004).
Skhul and Qafzeh in Israel) must make us 39. H. V. A. James, M. D. Petraglia, Curr. Anthropol. 46 77. I thank P. Forster, T. Kivisild, P. Endicott, P. Underhill,
aware that not all modern human dispersal (suppl.), S3 (2005). F. Balloux, C. Tyler-Smith, R. Foley, M. Lahr, M. Petraglia,
40. M. D. Petraglia, A. Alsharekh, Antiquity 77, 671 H. James, S. Jones, J. Koshy, L. Raddatz, S. Oppenheimer,
events were necessarily successful in either the (2003). S. Ambrose, A. Marks, C. Henshilwood, C. Marean,
long or short term, or in terms of their con- 41. S. A. Sali, The Upper Palaeolithic and Mesolithic Cultures O. Bar-Yosef, P. Van Peer, P. Vermeersch, I. Davidson,
tinuity in the subsequent human DNA record of Maharashtra (Deccan College, Poona, India, 1989). G. Barker, and T. Reynolds for discussion of points raised
(1, 2, 17, 29, 32). Even if the broad outlines of 42. M. Petraglia, personal communication. in the paper, and D. Kemp for assistance with the
43. S. U. Deraniyagala, The Prehistory of Sri Lanka: illustrations. Research funds were provided by the British
the modern human dispersal from Africa into An Ecological Perspective (Department of Archaeological Academy and Corpus Christi College, Cambridge.
Eurasia are now becoming much clearer, in Survey, Government of Sri Lanka, Colombo, Sri Lanka,
terms of both the genetic and archaeological 1992). 10.1126/science.1128402
800 11 AUGUST 2006 VOL 313 SCIENCE www.sciencemag.org