Genetics, Archaeology, and Holocene Hunter-Gatherers

Author(s): James F. O'Connell

Source: Proceedings of the National Academy of Sciences of the United States of America,
Vol. 96, No. 19, (Sep. 14, 1999), pp. 10562-10563
Published by: National Academy of Sciences
Stable URL: http://www.jstor.org/stable/48788
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Proc. Natl. Acad. Sci. USA
Vol. 96, pp. 10562-10563, September 1999
Commentary
Genetics, archaeology, and Holocer Lehunter-gatherers
James F. O'Connell*
Department of Anthropology, 270 South 14th Street East, Universityof Utah, Salt Lc ke City,UT 84112
Excoffier and Schneider's (1) analysis of mtDNA diversity in
a large sample of modern human populations yields results
consistent with those of previous work, indicating a series of
sharp expansions in human population size during the early
Upper Pleistocene [50-100 thousand years ago (KY)]. These
expansions perhaps originated first in Africa and then pro-
ceeded stepwise across Asia and the Americas and later into
Europe. Surprisingly, however, not all populations in Excoffier
and Schneider's (1) sample show evidence of these expansions;
the exceptions include several Native American and hunter-
gatherer groups. These are explained by appeal to recent or
recurrent population crashes or other processes likely to have
eliminated genetic evidence of earlier growth. Especially pro-
vocative is the suggestion that recent hunter-gatherer popula-
tions as a class may be smaller and more isolated than their
Pleistocene predecessors and thus more prone to demographic
upset. The utility of such populations as models for the analysis
of human demography in the Pleistocene is drawn into ques-
tion accordingly.
Genetic vs. Archaeological Evidence of Ancient Population
Growth. If Excoffier and Schneider (1) are right about early
Upper Pleistocene population increases, then the archaeolog-
ical record should offer some support, possibly including
indications of any behavioral changes that might have facili-
tated such growth, certainly by providing evidence of popu-
lation increases themselves (e.g., more sites, higher rates of
refuse deposition, and more colonization of areas previously
unoccupied).
As it happens, the anticipated matches are mixed at best;
major changes in the archaeological record generally trail the
dates proposed by Excoffier and Schneider (1) and others,
sometimes by tens of millennia. The correlation is best in
Africa, Europe, and parts of Asia, where the first appearance
of the Upper Paleolithic-complete with evidence of art,
formal burial, and other indications of modern human behav-
ioral capabilities-is generally dated at about 35-50 KY. These
dates overlap the more recent ends of the 95% confidence
intervals bracketing the proposed period of population growth,
although they fall short of many of the mean T values,
particularly in Africa (2). The same is true for Australia/New
Guinea, where the earliest archaeological dates are in the
range 40-60 KY (3), again overlapping the upper end of the
95% confidence interval but falling well short of three of four
mean r estimates cited for this region (76-97 KY). The
mismatch is most striking in the Americas. There, dates for two
indigenous populations showing what Excoffier and Schneider
(1) take to be clear indications of Pleistocene expansions are
55 and 59 KY (confidence intervals of 26-99 and 28-82 KY,
respectively); however, the earliest generally accepted archae-
ological dates are no greater than 13 KY (4, 5), Excoffier and
Schneider's (1) assertions about the existence and reliability of
30-35 KY dates notwithstanding (6).
These discrepancies may mean that (i) the accumulation of
recognizable archaeological evidence for population expan-
sions often trailed the expansions themselves by very substan-
tial lengths of time; (ii) the estimated mtDNA mutation rate is
PNAS is availableonline at www.pnas.org.
10 562
too low, making the genetically based dates for expansions too
early; or (iii) dates for population expansion in what are now
identified as non-African lineages may be unrelated to those
for the colonization of other continents. The first of these
possibilities seems unlikely, especially given the size of the
growth spurts posited by geneticists. Events of this magnitude
should have fairly clear and immediate archaeological conse-
quences. The idea that the mutation rate might be pegged too
low seems a stronger possibility. Excoffier and Schneider's (1)
reasons for rejecting this idea, notably that dates for mito-
chondrial Eve would have to be adjusted forward, are not
entirely compelling. The third alternative, that population
expansions might have begun well before continental coloni-
zations, seems quite likely, particularly for Australia/New
Guinea and the Americas. Both could have been occupied by
multiple groups that had already diversified elsewhere well
before entering either of these regions.
Genetic Diversity Not Explained by Stepwise Expansions.
Excoffier and Schneider's (1) suggestion that the absence of
evidence for Pleistocene population growth in some modern
groups is a reflection of recent demographic upset seems quite
plausible. The wonder is that such upsets have not disrupted
the genetic record of ancient demographic change more
thoroughly. Excoffier and Schneider (1) cite the catastrophic
impact of European colonization on certain Native American
populations. In fact, its effects were widespread on a conti-
nental scale, not only in the Americas (7, 8) but elsewhere (9).
Relatively recent climatic changes may also have had im-
portant demographic consequences, especially at the Pleisto-
cene/Holocene boundary (10-15 KY), when mean annual
temperatures worldwide rose and fell repeatedly over ranges
>5?C across periods of less than a century, sometimes less than
a decade (10, 11). It is not hard to imagine that human
populations everywhere suffered greatly as a result, reinforcing
the notion that the growth spurts indicated for many of those
included in the sample of Excoffier and Schneider (1) might
have occurred much later than many geneticists think and
again raising questions about the assumed mtDNA mutation
rate. Perhaps the idea that at least some were terminal (rather
than early Upper) Pleistocene phenomena is not far-fetched.
Recent Hunter-Gatherer Populations as Analogues for the
Pleistocene. The notion that historic and modern foragers, like
the !Kung, can be taken as Pleistocene relics has been chal-
lenged repeatedly by anthropologists over the past 3 decades,
and for good reasons. Most live in relatively unproductive
habitats, some of which were unoccupied before the end of the
last glaciation. Many are surrounded by agriculturists. Some
are embedded in large, state-level political systems of a kind
that have existed for no more than a few thousand years (12).
That said, it would be a mistake to infer, following Excoffier
and Schneider (1), that, as a class, historic and modern foragers
are genetically more isolated and that their effective popula-
tion sizes are thus necessarily smaller than those of their
Pleistocene predecessors. All historic and modern hunter-
The companionto this Commentarybegins on page 10597.
*To whom reprint requests should be addressed. E-mail: oconnell@
anthro.utah.edu.
 Commentary: O'Connell
gatherers have different, generally more complex extractive
technologies than did Pleistocene foragers. Most historic and
modern foragers exploit much broader arraysof resources, and
some have markedly higher local population densities, espe-
cially from the mid-Holocene onward. A few occupied very
productive habitats, notably in parts of the Americas (12-14).
Genetic, ethnographic, and archaeological data indicate that
those hunter-gatherers with agricultural neighbors interacted
reproductively with them over long periods, probably from the
time the groups first came into contact (15-17). Moreover, at
least some historic and modern foragers have moved repeat-
edly between farming and foraging lifestyles within the last few
millennia (17). Those without farming neighbors (and there
were many across all of Australia and over very large parts of
the Americas) had equally extensive contact with surrounding
hunter-gatherer groups (14, 18-20).
In short, it is hard to see recent hunter-gatherers overall as
either genetically more isolated or demographically more
fragile than those of the Pleistocene simply as a function of the
spread of agriculture. Abandoning them as a source of demo-
graphic insight on the distant past seems premature. It is better
to assess the utility of each population as a potential analogue
independently rather than to accept or reject them all cate-
gorically.
Excoffier and Schneider's (1) analysis is important in that it
helps move genetic research on recent human evolution away
from the "African Eve vs. multiregional origins" argument that
has dominated the literature over the last decade toward a
broader and, in many ways, more interesting set of questions.
The techniques they deploy clearly have enormous positive
potential. Fully realizing that potential will require active
collaboration between geneticists and their colleagues in the
historical sciences, especially anthropology.
1. Excoffier, L. & Schneider, S. (1999) Proc. Natl. Acad. Sci. USA
96, 10597-10602.
2. Klein, R. G. (1999) The Human Career: Human Biological and
Cultural Origins (Univ. Chicago Press, Chicago), 2nd Ed.
Proc. Natl. Acad. Sci. USA 96 (1999) 10563
3. O'Connell, J. F. & Allen, F. J. (1998) Evol. Anthropol. 6, 132-146.
4. Grayson, D. K. (1998) Science 282, 1425-1426.
5. Meltzer, D. J. (1993) Evol. Anthropol. 1, 157-169.
6. Meltzer, D. J., Adovasio, J. M. & Dillahay, T. D. (1994) Antiquity
68, 695-714.
7. Dobyns, H. F. (1983) Their Number Became Thinned: Native
American Population Dynamics in Eastern North America (Univ.
Tennessee Press, Knoxville, TN).
8. Ramenofsky, A. F. (1987) Vectors of Death: The Archaeology of
European Contact (Univ. New Mexico Press, Albuquerque, NM).
9. Butlin, N. G. (1983) Our OriginalAggression:Aboriginal Popula-
tions of Southeastern Australia 1788-1850 (Allen & Unwin,
Sydney).
10. Taylor, K. C., Lamorey, G. W., Doyle, G. A., Alley, R. B.,
Grootes, P. M., Mayewski, P. A., White, J. W. C. & Barlow, L. K.
(1993) Nature (London) 361, 432-435.
11. Taylor, K. C., Mayewski, P. A., Alley, R. B., Brook, E. J., Gow,
A. J., Grootes, P. M., Meese, D. A., Saltzman, E. S., Severinghaus,
J. P., Twickler, M. S., et al. (1997) Science 278, 825-827.
12. Kelly, R. L. (1995) The Foraging Spectrum: Diversity in Hunter-
GathererLifeways (Smithsonian Inst. Press, Washington, DC).
13. Price, T. D. & Brown, J. A., eds. (1985) Prehistoric Hunter-
Gatherers: The Emergence of Cultural Complexity (Academic,
Orlando, FL).
14. Ames, K. M. & Maschner, H. D. G. (1999) Peoples of the
Northwest Coast: Their Archaeology and Prehistory (Thames &
Hudson, London).
15. Schrire, C., ed. (1984) Past and Present in Hunter-GathererStudies
(Academic, Orlando, FL).
16. Yellen, J. E. & Harpending, H. C. (1972) World Archaeol. 4,
244-253.
17. Barnard, A. (1992) Hunters and Herders of Southern Africa: A
Comparative Ethnography of the Khoisan Peoples (Cambridge
Univ. Press, Cambridge, U.K.).
18. Kirk, R. L. (1981) Aboriginal Man Adapting: The Human Biology
of Australian Aborigines (Clarendon, Oxford).
19. Mulvaney, D. J. (1976) in Tribes and Boundaries in Australia, ed.
Peterson, N. (Aust. Inst. Aboriginal Stud., Canberra, Australia).
20. Heizer, R. F. (1978) Handbook of North American Indians
(Smithsonian Inst. Press, Washington, DC), Vol. 8.