rapid expansion in these specific mito-
chondrial types amongst the existing Afri-
can populations, remains perhaps more
debatable. But in either case, it is clear
that some significant demographic or cul-
tural factors must have promoted these
lineage expansions at roughly the same
time as the mtDNA mismatch analyses
point to a rapid increase in total popula-
Fig. 1. mtDNA ‘‘mismatch’’ distributions of tion numbers from some localized geo-
present-day African, Asian, and European popula- graphical source. A similar expansion in
tions, showing the frequency distribution of differ- African populations has also been claimed
ences between pairs of individuals in the three pop- from some studies of DNA microsatellite
ulations. The modes of the three distributions clearly
data, although with less specific age esti-
reﬂect a much earlier demographic expansion of Af-
rican populations (ca. 80,000 B.P.), than those in Asia
(ca. 60,000 B.P.) and Europe (ca. 40,000 B.P.) (23–25).
The central question is what could have
ern African populations by Watson, For- caused this apparently dramatic expansion
ster, Salas, Kivisild, Macaulay, and others in African populations 60,000–80,000
(2, 8, 9, 26–28). Once again, the precise B.P., and it is here that recent archaeolog- Fig. 3. Map of archaeological sites and early
timing of these lineage expansions de- ical research in southern and central Af- anatomically modern human remains in Africa and
pends on the assumed mutation rate of rica becomes central to the interpretation Israel, referred to in text.
mtDNA, but, in all of these studies, there of the demographic data. The most rele-
is evidence for what Forster and Mat- vant evidence at present comes from a sively shaped bone tools, apparently used
sumura (28) have recently described as a number of sites located close to the south- as both tips of throwing spears and
‘‘remarkable expansion’’ of the distinctive ern tip of Africa in Cape Province, most sharply pointed awls for skin working (36,
L2 and L3 mitochondrial lineages dating notably from Blombos Cave and Klasies 37); new forms of carefully shaped stone
broadly to between ca. 80,000 and 60,000 River on the southern coast and those of inserts, probably used as tips and barbs of
B.P. (2, 8, 9, 26–28) (Fig. 2). As in the Boomplaas Cave and Diepkloof, further either hafted throwing spears or conceiv-
case of the mismatch analyses, the evi- to the north and west (29–40) (Fig. 3). ably wooden arrows (30–32, 34, 51); large
dence points to an expansion centered These are backed up by a number of numbers of perforated estuarine shells,
initially in one small area of Africa (most rather less well documented sites in east- evidently used as personal ornaments of
probably in eastern or southern Africa) ern and central Africa (34, 41–43). The some kind (39); and large quantities of
followed by an expansion to other regions, general time range of these sites is that of imported red ochre, including two pieces
the African Middle Stone Age (MSA) from the Blombos cave with carefully in-
apparently reaching western Africa by at
extending from 250,000 to 40,000 B.P., cised and relatively complex geometrical
least 30,000–40,000 B.P., and perhaps designs on their surfaces (38). These de-
across the mouth of the Red Sea to the and coinciding broadly with the Middle
Palaeolithic (or Mousterian) periods in signs represent the earliest unambiguous
adjacent parts of southern Asia by forms of abstract ‘‘art’’ so far recorded
60,000–65,000 B.P. (2, 8, 9, 28). Europe and Asia (44, 45). But the rele-
vant evidence from the so-called ‘‘Still (Figs. 4 and 5). Equally significant in
Whether this dispersal of the L2 and L3 these sites is the evidence for the large-
Bay’’ levels in the Blombos Cave and the
lineages reflects an actual dispersal of dis- scale distribution or exchange of both
ensuing ‘‘Howiesons Poort’’ levels at Kla-
crete human populations, or simply a high-quality stone for tool production and
sies River, Boomplaas, and Diepkloof, can
be dated specifically to the later stages of the recently discovered shell beads from
the MSA, between ca. 75,000 and 55,000 the Blombos cave, in both cases either
B.P. (35, 46, 47). transported or traded over distances of at
least 20–30 km (31, 39). All of these fea-
Although the archaeological assem-
tures show a striking resemblance to those
blages from these sites have traditionally
which characterize fully modern or ‘‘Up-
been attributed to the MSA, they reveal a
per Palaeolithic’’ cultures in Europe and
number of radical technological and cul- western Asia, which first appeared with
tural features that collectively contrast the initial arrival of anatomically and be-
sharply with those of the earlier African haviorally modern populations at
MSA sites, and which show many resem- 45,000–50,000 B.P., i.e., some 20,000
blances to those that appear in Europe years later than their appearance in the
and western Asia with the arrival of the African sites (17, 45, 48–50). As Hen-
first anatomically and genetically modern shilwood (35) has recently commented,
populations at 45,000–50,000 B.P., the the combination of these behavioral inno-
period of the so-called ‘‘Upper Palaeo- vations in the Still Bay and succeeding
lithic revolution’’ (17, 45, 48–50). These Howiesons Poort levels at these South
assemblages include, for example, new African sites seems to reflect ‘‘a dynamic
patterns of blade technology, produced by period of diverse technological behavior
means of ‘‘soft hammer’’ techniques of not previously seen in the African Middle
Fig. 2. Inferred patterns of geographical dis-
persal of the L2 and L3 mtDNA lineages in Africa
flaking (29–32, 51); new forms of both Stone Age.’’
between ca. 80,000 and 60,000 B.P., according to specialized skin working tools (end-scrap-
Forster (2). Later dispersals of the M, N, and R ers) and tools for the controlled shaping Population Expansion
lineages into Asia and Europe after 65,000 B.P. of bone and wooden artefacts (so-called The critical importance of these new ar-
derive from the L3 lineage. burin forms) (32, 35); a range of exten- chaeological discoveries is that they may
9382 www.pnas.org cgi doi 10.1073 pnas.0510792103 Mellars
groups (31, 34). The second and poten- systematic exploitation of marine fish, and
tially equally important suggestion, which perhaps sea birds, as parts of the human
has been mooted by Deacon (29, 31), is food supply. Finally, Deacon, Ambrose,
that the dense accumulations of burnt and others (31, 35, 42, 43) have argued
plant remains in the Howiesons Poort that the large-scale movements of high-
levels at Klasies River (together with quality stone and imported shell orna-
identifiable remains of root crops, such as ments recorded from these sites may
Watsonia, in the later MSA levels at the reflect increased trading and exchange
Strathalan B site, further to the north) networks between adjacent human groups,
could reflect either the increased use of which could have acted as a further criti-
these particular plant resources or even cal mechanism to ensure regular access
the deliberate burning of the local fynbos and distribution of essential food supplies,
vegetation, which has been shown to in- especially during seasonal or other epi-
crease the annual productivity of these sodes of food scarcity.
root crops by between five- and ten-fold Clearly, all of these possibilities will
(31). For the present, the latter suggestion require further analysis and testing in the
remains speculative, but, if this were to be course of future research. But the implica-
supported by further research, one could tion seems clear that many of the behav-
see this effectively as an early form of ioral innovations reflected in the southern
plant food management strategies, poten- African archaeological records between
tially analogous to those used in later ca. 80,000 and 60,000 B.P. could have led
Mesolithic and early agricultural commu- to a substantial increase in the carrying
Fig. 4. Stone tools from the MSA Howiesons Poort
nities, or in the recently reported 26,000- capacity of the environment for human
levels at Klasies River (South Africa) dated to ca.
65,000 B.P., showing closely similar forms of blades,
year-old processing of seed remains from populations and, accordingly, to a major
end scrapers, burins, and small, hafted segment the Ohalo II site in Israel (52). A third expansion in human population numbers
forms to those found in European and Asian Upper suggestion advanced by Henshilwood (35, and densities. Even allowing for the im-
Palaeolithic sites from ca. 45,000 B.P. onwards (32). 36) is that the Still Bay levels at Blombos precisions in current DNA dating esti-
cave may provide evidence for the first mates, the apparent coincidence between
provide the explanation for the major ex- these major behavioral changes and the
pansion in African populations, which is estimated timing of the population expan-
reflected so clearly in the recent mtDNA sions reflected strongly in both the
evidence, dated broadly to between 80,000 mtDNA mismatch and lineage-analysis
data seems hard to ignore. It should be
and 60,000 B.P. The precise cultural and
emphasized that there is no necessary im-
demographic mechanisms that underlay
plication that population numbers in Af-
the population expansion inevitably re-
rica as a whole increased dramatically at
main more hypothetical. At least four as-
this time. Indeed, it could be that total
pects of the archaeological data, however,
population numbers in Africa decreased
could be significant in this context. The
significantly at this time, owing to the on-
first is that the character of the artefacts set of extremely dry conditions in many
recovered from both the Blombos Cave parts of Africa between ca. 60,000 and
and the Howiesons Poort levels at Klasies 30,000 B.P. (31, 44). The point is simply
River and elsewhere would appear to re- that increased levels of technological effi-
flect the emergence of more complex ciency and economic productivity in one
forms of hunting equipment, apparently small region of Africa could have allowed
involving the construction of several dif- a rapid expansion of these populations to
ferent forms of hunting weapons (i.e., the other regions and an associated competi-
sharply pointed bone spear heads and the tive replacement (or absorption) of the
bifacial leaf-point forms from the Blom- earlier, technologically less ‘‘advanced,’’
bos cave, and the appearance of compos- populations in these regions (2, 16, 23,
ite, multiple-component hafted weapons 53, 54).
in the Howiesons Poort levels at Klasies Any attempt to define the precise point
River and other sites) (31, 34–37) (Fig. 4). of origin of these behavioral innovations,
The possibility has been suggested that and the associated demographic expansion
some of these forms could well have event, immediately encounters the relative
served as the tips and barbs of wooden sparsity of well documented archaeologi-
arrows, based on comparisons with similar cal sites in many regions of subSaharan
artefacts recovered from both later Afri- Africa, especially in the more central and
can Stone Age sites and much later Meso- eastern areas of Africa, which are poten-
lithic contexts in Europe (34, 40). Even tially crucial to the current debates over
without inferring the use of archery Fig. 5. Fragments of red ochre incised with a modern human origins (31, 34). Clearly,
equipment, however, it is reasonable to complex geometrical design (A–D), and a series of we must be aware of falling into the obvi-
assume that the introduction of more ef- deliberately perforated shells of Nassarius krauss-
ous trap of assuming these developments
ianus (E) from the MSA levels of the Blombos Cave
fective hunting weapons would have sub- must have occurred initially within South
(South Africa), dated to ca. 75,000 B.P. (38, 39).
stantially increased the efficiency and pro- [A–D reproduced with permission from Hen- Africa, simply because this area is where
ductivity of hunting activities and, shilwood et al. (38) (Copyright 2002, AAAS). E re- the relevant archaeological evidence is at
therefore, the overall productivity of the produced with permission from Henshilwood et al. present most fully investigated and best
food resources available to the human (104) (Copyright 2004, AAAS).] documented (i.e., the ‘‘drunk looking for
Mellars PNAS June 20, 2006 vol. 103 no. 25 9383
keys under the street lamp’’ syndrome!). in human technology, subsistence, settle-
In this context, it should be recalled that ment patterns, and associated patterns of
industries conforming closely to the South social and even symbolic communication
African Howiesons Poort variations are as a fairly direct response to the new envi-
well represented over large areas of cen- ronmental challenges that emerged at this
tral and southern Africa (to the south of time (53, 54, 61, 62). Significantly, all
the Zambezi) and apparently extending these major environmental changes fall
northwards into parts of East Africa— within the time range of ca. 80,000–70,000
such as at the site of Mumba in Tanzania B.P., precisely the time when the archaeo-
(34, 41) and the recently excavated site of logical evidence indicates that technologi-
Norikiushan in Kenya (S. Ambrose, per- cal and other behavioral changes were
sonal communication) 4,000 km to the occurring most rapidly.
north of the South African sites. The
main problem at present lies in the accu- Human Cognitive Evolution
rate dating of these sites in relation to the Even if we accept that the pattern of be-
South African localities (34). On present havioral changes in southern Africa can
evidence, it is impossible to exclude the be explained more parsimoniously in
possibility that the Howiesons Poort tech- terms of adaptive environmental processes
nologies, or indeed those of the preceding than by changes in human cognitive ca-
Still Bay, could have emerged in certain pacities, we cannot escape the evidence
parts of, say, eastern or central Africa, for significant changes in at least some
before they subsequently appeared in the aspects of human cognitive behavior asso-
South African sites. In this case, the de- ciated broadly with the emergence of our
velopments in South Africa could be seen own species (63–68). One aspect of the
more as a reflection of events in other current evidence that is potentially highly
parts of Africa than their initial point of informative in this context lies in the evi-
origin. But, in any event, the sheer scale dence for a precocious and apparently
of the geographical distribution of the Fig. 6. Summary of the model proposed here for short-lived expansion of anatomically
Howiesons Poort-like technologies could modern human origins and dispersal from Africa. modern populations from northern Africa
be seen as a further potential reflection of into the immediately adjacent areas of
a major episode of population dispersal southwest Asia at 110,000–90,000 B.P.
logical mutation (although, according to (1, 69–72). This expansion is best re-
within subSaharan Africa centered
the model advanced here at 80,000 B.P. flected in the large samples of typically (if
broadly within the time range from ca.
and not at ca. 40,000–50,000 B.P., as relatively robust and variable) anatomi-
70,000 to 55,000 B.P. (31, 34, 35). It is
Klein himself has suggested). Or alterna- cally modern skeletal remains from the
equally tempting to suggest that it was
tively (and more prosaically), we could two sites of Skhul and Qafzeh in northern
precisely this new, integrated complex of
look for an interpretation in terms of Israel (Fig. 3). Three features of these
so-called modern behavioral features
some major shift in the adaptive and se- finds are especially significant. The first is
embodied in the Howiesons Poort and
lective pressures to which the human that at least two of the skeletons in these
preceding Still Bay technologies that led
populations were subjected, perhaps pre- sites occurred in the form of clearly cere-
directly to the widespread geographical
cipitated by some major episode of cli- monial or ritualistic burials, associated
expansion of the southern African popula-
matic and environmental change. In this with seemingly unmistakably intentional
tions not only to other areas of Africa (as
context, the obvious candidate would be grave offerings (a large deer antler lying
reflected in the widespread dispersal of
the sharp oscillations between wetter and directly on top of one of the Qafzeh skel-
the L2 and L3 mitochondrial lineages; see
drier climatic conditions that marked the etons and a complete boar’s jaw said to
Fig. 2) but also to the adjacent areas of
transition from oxygen isotope stage 5 to be ‘‘clasped in the arms’’ of one of the
Asia and Europe, sometime after 70,000
stage 4, as reflected in the deep-sea core burials at Skhul) (72–75) (see Fig. 7). Sec-
B.P. (1, 2, 8, 16, 17, 42) (Fig. 6).†
and ice-core climatic records (56). In sub- ondly, that, at least in the case of the
The Mechanisms of Behavioral Change Saharan Africa, there is evidence that this Qafzeh burials, the remains were associ-
transition resulted in changes in annual ated with a number of deliberately perfo-
The pivotal question, of course, is what
rainfall by up to 50% (57). To groups oc- rated seashell ornaments, together with
caused these radical changes in the tech-
cupying the more arid regions of Africa large quantities of used and apparently
nology, economy, and social patterns of
(especially around the margins of the heat-treated fragments of red ochre, al-
African groups 80,000–70,000 B.P.?
Kalahari and Sahara deserts), the impact most certainly used as coloring pigments
Here we have two fairly stark alternatives.
of these climatic changes on all aspects of (76, 77). And, thirdly, that, despite these
First, we could suggest, as Klein (44, 55)
human economic, technological, and so- clearly ‘‘symbolic’’ aspects of the archaeo-
has done, that the emergence of distinc-
cial adaptations could have been dramatic, logical material, the stone tool assem-
tively modern patterns of culture and
as Deacon, Ambrose, and others (29, 31, blages found in association with both the
technology was due to a sudden change in
42, 43) have emphasized. A further poten- Skhul and Qafzeh remains were of typi-
the cognitive capacities of the populations
tially significant factor could have been cally Middle Palaeolithic or MSA in form,
involved, entailing some form of neuro-
the climatic and associated environmental without any trace of the distinctively mod-
effects of the Mount Toba volcanic ‘‘su- ern or Upper Palaeolithic technological
†Note that claims for a reemergence of MSA-like technol- pereruption’’ in Sumatra, dated to features recorded at the later African
ogies after the Howiesons Poort industries in South Africa 73,000 B.P., as Ambrose (58) has argued MSA sites of Klasies River, Blombos, and
(31) are not directly relevant to this model, because it is very effectively [but see Oppenheimer elsewhere (71, 72).
likely that by this time (ca. 50,000 –55,000 B.P.) the initial
dispersal from Africa had already taken place (8, 9, 28).
(59) and Gathorne-Hardy & Harcourt- The clear implication of these finds is
Exactly what these post-Howiesons Poort MSA industries Smith (60) for an opposing view]. It that, whilst the human populations repre-
represent remains to be clariﬁed. would, in short, be possible to see changes sented at Skhul and Qafzeh were essen-
9384 www.pnas.org cgi doi 10.1073 pnas.0510792103 Mellars
two early anatomically modern skulls re- complexity of technological and other be-
cently discovered at Herto in Ethiopia, havioral patterns for which the necessary
dated to 160,000 B.P., and again associ- cognitive potentials had already long ex-
ated with characteristically archaic MSA isted (68, 82, 84, 92)? One thing, however,
stone tool technology (6, 79). is certain: If the evolutionary trajectories
If so, what, if anything, might this evi- of the Eurasian Neanderthals and the Af-
dence tell us tell us about the patterns of rican ancestors of modern populations
human cognitive and neurological evolu- had been separate over a span of at least
tion associated with the emergence of 300,000 years [as all of the current genetic
fully anatomically and genetically modern and skeletal evidence suggests (1, 12, 78,
populations? If explicit symbolism is ac- 93)], then the possibility of some signifi-
cepted as an index of essentially modern cant changes in human neurological and
cognitive capacities and with associated cognitive capacities over this time range
patterns of essentially modern, complex can in no way be ruled out. Even if the
language [as most archaeologists and cognition of Neanderthals and other
palaeoanthropologists tend to assume (30, archaic populations was not ‘‘inferior’’
63, 66, 68, 80–87)], then these capacities to that of modern humans, it could have
were clearly in place by at least 100,000– been significantly different (66, 67,
150,000 B.P. and could well have emerged 80–83).
in direct association with the evolution of
Fig. 7. Burial of an anatomically modern human
skeleton at the Qafzeh Cave (Israel), accompanied
anatomically and genetically modern pop- The Out of Africa Diaspora
by a large deer antler and dated to ca. 90,000 – ulations at this time. Viewed in these The final, and most controversial, issue at
100,000 B.P. (73, 74). terms, the subsequent elaboration of these present is exactly when and how these
symbolic patterns and the emergence of a anatomically and genetically modern pop-
range of new technological, economic, and ulations first spread from Africa to other
tially modern in both anatomical terms
social patterns reflected in the archaeo- parts of Asia and Europe. Here there are
and in terms of clearly symbolic behav-
logical evidence from Blombos, Klasies two main possibilities. The first is that the
ioral patterns, the levels of technology
River, and elsewhere, could be seen sim- initial expansion occurred via North Af-
associated with these populations were
ply as a gradual working out of these new rica and the Nile valley, with subsequent
still of strictly archaic, Middle Palaeolithic cognitive capacities under the stimulus of
form (71, 72). Viewed in these terms, it is dispersals to both the west into Europe
various kinds of environmental, demo-
equally interesting that the early incursion and to the east into Asia (69–71, 78, 94,
graphic, or social pressures, in much the
of these anatomically modern populations 95). The second is that the initial dispersal
same way as that reflected in the later
into southwest Asia seems to have been a was from Ethiopia, across the mouth of
emergence of fully agricultural communi-
very localized and short-lived event, ap- the Red Sea, and then either northward
ties (53, 54, 61, 68, 88).
parently confined to this southwest Asian through Arabia or eastward along the
The alternative, of course, would be
region, and followed by a reestablishment south Asian coastline to Australasia—the
to visualize the trajectory of human cog-
of the earlier Neanderthal populations nitive evolution as an inherently more so-called ‘‘southern’’ or ‘‘coastal’’ route
within these regions from at least 70,000 complex process, involving potentially a (28, 69, 70, 96). The strongest evidence at
B.P. onwards, as reflected by the typically series of successive and cumulative present for the second hypothesis is pro-
Neanderthal remains recovered from the changes in brain capacities, dependent vided by the mtDNA lineage-analysis
later Mousterian levels at the Kebara on a succession of genetic mutations patterns. These point strongly to the con-
cave, Tabun, Amud and Shanidar (1, 71, affecting various aspects of brain func- clusion that there was only a single (suc-
72, 78). In other words, it would seem that tion and organization (63–67, 89, 90). cessful) dispersal event out of Africa,
whatever the intellectual and symbolic Recent studies of the Microcephalin and represented exclusively by members of the
capacities of these early anatomically FOXP2 genes (63, 64) have now effec- L3 lineage and probably carried by a rela-
modern populations, their levels of tech- tively demonstrated the possibility of tively small number of at most a few hun-
nological and socioeconomic organization such mutations, potentially at various dred colonists (2, 8, 28, 97). This lineage
were not sufficient to withstand competi- points since the emergence of geneti- rapidly diversified into the derivative M,
tion from the long-established Neander- cally modern humans. Clearly, if there N, and R lineages, which are particularly
thal populations of Eurasia during the had been a further genetic mutation in- well represented in modern Asian popula-
later (and colder) stages of the Middle volving cognitive capacities 80,000 tions and which are estimated to have ar-
Palaeolithic sequence (71, 72, 78). B.P., this could provide a further poten- rived and diversified further in southern
tial explanation for the emergence of Asia by at least 50,000 B.P. and possibly
Mosaic Evolution significantly new patterns of technology, as early as 65,000 B.P. in Malaysia and the
The obvious and seemingly inescapable social organization, and symbolic expres- Andaman islands (8, 9, 28, 97). A similar
conclusion is that the patterns of cultural sion reflected in the archaeological evi- conclusion has been drawn from recent
and technological development associated dence from the African sites. studies of the Y chromosome evidence
with the evolution of fully modern popula- The problem of adequately testing these (97). This evidence would also conform
tions were strongly mosaic in character, speculations against hard archaeological well with the clear peak in the mtDNA
with the emergence of several explicitly data is, of course, one of the notorious distributions of Asian populations, dated
symbolic aspects of culture apparently dilemmas in studies of human cognitive broadly to 60,000 B.P. (23–25) (Fig. 1).
preceding any major change in either the evolution, epitomized by Renfrew’s (91) This model, of course, would mean that
stone-tool or bone-tool components of the notion of the ‘‘Sapient paradox.’’ In other the subsequent dispersals of anatomically
associated technologies (40). In Africa words, how do we formulate plausible and behaviorally modern populations into
itself, there may be further evidence for archaeological tests for the emergence of southwest Asia and Europe must have
this symbolic behavior in the indications new behavioral capacities, as opposed to reached these areas substantially later, via
of apparently ritualistic treatment of the the gradual elaboration and increasing western or central Asia (2, 8, 97).
Mellars PNAS June 20, 2006 vol. 103 no. 25 9385
The main problem posed by this sce- (98–100). But clearly the spotlight is I thank P. Forster, S. Matsumra, C. Stringer,
nario at present lies in the sparsity of now directed strongly onto southern H. Harpending, A. Rogers, T. Kivisild, P.
well documented and well dated archae- Asia to secure more direct evidence for Underhill, C. Marean, P. Endicott, A. Brooks,
ological evidence for the early modern this hypothetical early dispersal route S. Ambrose, G. Sampson, O. Bar Yosef, C.
Henshilwood, J.-J. Hublin, and M. Petraglia for
human colonization of Asia prior to ca. (101, 102). Future discoveries in both
discussions of points raised in the paper. I also
45,000 B.P., when we know that early mitochondrial and Y chromosome DNA thank D. Kemp for assistance with the illustra-
colonists had reached parts of northern research and, above all, archaeology, are tions, P. Forster for Fig. 2, and C. Henshilwood
and southern Australia, best represented awaited to provide the crucial tests for for Fig. 5. Research funds were provided by the
by the archaeological and skeletal finds this hypothesis of the origins and dis- British Academy and Corpus Christi College
from Lake Mungo in New South Wales persal of our own species (103). (Cambridge University, U.K.).
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