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Where River Meets Sea
The Southern-Dispersal Hypothesis
A Parsimonious Model for Homo sapiens
Colonization of the Indian Ocean Rim and Sahul
David Bulbeck
For approximately the past decade, the Out-of-Africa or re-
placement theory has enjoyed the status of the new orthodoxy
on the origins of H. sapiens. It is certainly compatible with
the fossil record (Stringer and McKie 1996; Cameron and
Groves 2004) and is strongly supported by the mitochondrial
DNA and Y-chromosome evidence (Oppenheimer 2003; Har-
ding and Liu 2005). As emphasized by Oppenheimer (2004,
175–84), Macaulay et al. (2005 a ), and Thangarej et al. (2005),
the rakelike structure of deeply rooted mtDNA lineages (hap-
lotypes M and N) right along South Asia, Southeast Asia, and
Australia–New Guinea indicates a Late Pleistocene dispersal
event that was rapid on an archaeological time scale.
The most creditable opposition comes from geneticists
working with nuclear DNA, who reconstruct time depths for
many non-African polymorphisms on the order of hundreds
of thousands of years. Commenting on the three Science ar-
ticles referenced above, Harpending and Eswaran (2005)
briefly summarized the misgivings of human nuclear genet-
icists about the Out-of-Africa theory. In response, Macaulay
et al. ( 2005 b ) pointed to methodological problems with the
time depths estimated by nuclear geneticists and noted that
our article (Macaulay et al. 2005 a ) had not touched on
whether H. sapiens might have incorporated relics of other
populations during its Out-of-Africa expansion. I leave ad-
judication on the genetic evidence to my qualified colleagues
(Richards et al. 2006) and address this article to readers not
entirely opposed to the Out-of-Africa theory.
The rapid-southern-dispersal hypothesis could possibly be
critiqued in terms of the absence of a rationale for the col-
onists’ having kept moving ever eastward. Spencer Wells (re-
ported in Shreeve 2006, 68) suggests that “it was less of a
journey and more like walking a little further down the beach
to get away from the crowd.” This idea, however, raises the
questions why individual colonists had wished to escape their
social milieu and how they had survived away from their
group. The colonization model presented here will resolve
such questions. It is inspired more by the mtDNA evidence
than by other considerations, though its relevance for ar-
chaeology will be addressed.
School of Archaeology and Anthropology, The Australian
National University, ACT 0200, Australia (david.bulbeck@
anu.edu.au). 31 X 06
Recent genetic research suggests an expansion along the trop-
ical coastline of the Indian Ocean, between 75,000 and 60,000
years ago, of the population which included the ancestors of
all of the non-African human mitochondrial DNA lineages
known today. In view of the arid sections along this coastal
stretch, irregularly punctuated by resource-rich estuaries, and
the crossings over open sea during the last leg to Australia/
New Guinea, this expansion would necessarily have involved
the features of watercraft, portage of potable water, and ad-
aptation to estuaries. These features could well have been the
cultural basis for the rapid tropical dispersal of Homo sapiens
out of Africa to Australia/New Guinea.
The Out-of-Africa theory proposes a Late Pleistocene expan-
sion of Homo sapiens from its African cradle into Eurasia.
Proponents of the theory have disagreed on whether there
had been a southern dispersal along the coastal belt of Arabia
and southern Asia, a more inland dispersal through the Mid-
dle East, or both (Kingdon 1993; Lahr and Foley 1994; Klein
1999; Oppenheimer 2003, 2004; Mellars 2006). The southern-
dispersal hypothesis is strengthened by mtDNA research
which suggests a rapid eastward migration of Homo sapiens
along the northern rim of the Indian Ocean, starting from
East Africa at around 75,000 years ago (Macaulay et al. 2005 a ;
Thangarej et al. 2005). As Forster and Matsumura (2005)
remark, this hypothesis has profound implications for un-
derstanding when early colonists to Wallacea and Sahul staged
the necessary water crossings from Sunda (fig. 1). In this
contribution I propose a model, based on an estuarine ad-
aptation, to account for the average rate of 0.7 to 4 km per
year calculated by Macaulay et al. (2005 a ) for the eastward
dispersal of H. sapiens. It is neither a “push” nor a “pull”
model and does not presume of the colonists any grand vision
of occupying and populating unknown terrain. Instead, it
proposes a tradition of seeking out suitable estuarine habitat
as the main impetus for the migratory movement.
Colonization along the Northern Rim of
the Indian Ocean
Well before the Out-of-Africa theory took root, Sauer (1963,
311) had proposed that “the dispersal of early man took place
most readily by following along the seashore.” Sauer pointed
out that coastally adapted populations would have had access
to a continuous string of familiar habitats little affected by
seasonal changes to the climate and biota. With regard to arid
landmasses, Sauer argued that desert shores are richer in food
resources than the adjacent hinterland and that even here
potable water could be found in seeps at low tides. “Coastwise
2007 by The Wenner-Gren Foundation for Anthropological Research.
All rights reserved. 0011-3204/2007/4802-0007$10.00
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Current Anthropology Volume 48, Number 2, April 2007
Figure 1. Proposed Out-of-Africa dispersal routes and sites mentioned
in the text.
there was scarcely a barrier to the spread of early man through
tropical and subtropical latitudes” (p. 312).
Sauer’s line of thought was further developed by Kingdon’s
(1993) attempt to relate the Out-of-Africa theory to mam-
malian dispersal routes. Kingdon suggested that Homo sapiens
strandloopers had essentially followed the marine shoreline
(or dugong–flying fox) route from the Bay of Bengal to Sahul
in one direction and along South Asia to East Africa in the
other (fig. 1). He noted the similarity in resources between
tropical rivers and tropical seas, especially at river mouths, in
mangrove swamps, and on the foreshore. He also remarked
that watercraft would have opened up reefs and marine shal-
lows as niches for human foragers. As for the original mi-
gration of Homo sapiens from Africa to the Bay of Bengal,
Kingdon hypothesized that these emigrants had taken the
tropical Indian or monkey-porcupine route. He dated this
event to 80,000–115,000 years ago on the basis of the dates
then available for anatomically modern skeletons from Skhul
and Qafzeh in the Levant (Kingdon 1993, 77–80, 120–22,
135–36).
Lahr and Foley (1994) foreshadowed the southern-dispersal
hypothesis in their model of two movements by Homo sapiens
out of Africa: a lesser migration along the northern rim of
the Indian Ocean and a subsequent main migration through
Asia Minor. Oppenheimer (2003) reversed the order and im-
portance of these dispersals. He associated the Skhul/Qafzeh
people with a failed early exodus from Africa, as their age of
90,000–120,000 years antedates the estimated age of
70,000–85,000 years for the common ancestor of all non-
African mtDNA. He also noted that at around 80,000 years
ago the overland route out of Africa would have been con-
fronted by continuous desert. In contrast, the southerly route
across the Hormuz Strait would have been assisted by patches
of coastal scrub along East Africa and the southern Arabian
Peninsula. Observing that the Eritrean site of Abdul docu-
mented the human exploitation of reef resources by 125,000
years ago, Oppenheimer (2003) proposed a dispersal of
“beachcombers” from East Africa to Sahul along the northern
rim of the Indian Ocean (fig. 1). Macaulay et al. (2005 a )
subsequently proposed a slightly later chronology, beginning
in the Horn of Africa at around 75,000 years ago and reaching
Sahul at around 60,000 years ago.
The final leg to Sahul would clearly have involved watercraft
capable of voyages of 50–100 kilometers (Mulvaney and Kam-
minga 1999, 109–10). While the colonization of Australia by
at least
years ago is clear (Gillespie 2002), the Wal-
50,000
lacea archaeological record is more complex. Flores, with its
highly distinctive record of hinterland habitation from the
Early Pleistocene till 12,000 years ago (Morwood et al. 2005;
Brumm et al. 2006), stands apart from the other islands of
Wallacea, whose well-documented archaeological time depth
is consistently 35,000 years or less (Bellwood 2001; Bulbeck
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317
2004). Irrespective of the enigma surrounding the precocious
settlement of Flores, there is minimal evidence for an Aus-
tralasian maritime capacity that could have reached Australia
before 60,000 years ago (Gillespie 2002; O’Connor and Chap-
pell 2003) or a widespread colonization of Wallacea before
35,000 years ago.
The arrival of Homo sapiens in Australasia at around 60,000
years ago, as indicated by the genetic evidence (see also van
Holst Pellekaan et al. 2006), is the likely cause for the new-
found capacity to colonize Australia. Given the dating im-
precisions, this could have occurred as much as 10,000 years
after Homo sapiens had reached Sunda—a long period of time
for the local development of a maritime capacity. By the same
token, the crossing to Sahul could have occurred instanta-
neously (in archaeological terms), as asserted by the rapid-
southern-dispersal hypothesis. In that case it would be likely
the beachcombers already had boats when they arrived rather
than fortuitously inventing them on their arrival. This is en-
tirely reasonable in view of the archaeological evidence from
East Africa of fully modern capacities in artifact manufacture
and exploitation of littoral and reef resources by 125,000 years
ago (Walter et al. 2000). Thus, acceptance of the rapid-south-
ern-dispersal hypothesis would strongly suggest watercraft
assistance.
enclosed coastal embayment where fresh water and seawater
meet and mix,” as the point of origin for successful lengthy
forays.
All of the advantages mentioned by Sauer (1963) and King-
don (1993) for coastal foragers would have been optimized
in the Indian Ocean’s estuaries. Timber (for boat building)
and potable water would have been available, even if requiring
some upstream travel. The occurrence of coral reefs, sea-grass
beds, and/or mangrove belts along the northern rim of the
Indian Ocean (Sheppard 2000) would have supported a sim-
ilar range and abundance of marine and littoral resources
along the entire route. This continuity of familiar resources
would also have prevailed at river mouths and stream outlets,
with many of the same or closely related organisms as are
found at other coastal locations (Nybakken 2001, 335–38).
Finally, watercraft would have provided protection against
saltwater crocodiles and other potential dangers.
Given their rich resources, estuaries would have supported
healthy population growth, conveying an impression of
crowded conditions even if flanked by stretches of vacant land.
This would have stimulated the search for new estuarine hab-
itats, which, to the experienced, advertise their presence out
to sea. Colonists towards the front of the wave would have
known how to travel back to their parent community, when-
ever conditions were ripe, producing a chain of interlinked
estuarine settlements (cf. Moore 2001). In summary, an es-
tuarine specialization would have been the ideal adaptation
to carry colonizing groups out of Africa past the Arabian
Peninsula—a demanding section of the route in terms of
drinking water (Oppenheimer 2004, 68; Petraglia and Al-
sharekh 2003, 681)—to South Asia and onward to Sunda.
Virtually the entire route is tropical (fig. 1), minimizing bar-
riers against the passage of individuals maladapted to cooler
conditions.
The questions raised earlier with respect to Wells’s suggestion
of “beachcomber creep” are readily resolved by the model of
estuarine-focused paddlers. The model specifically invokes a
chain of linked estuarine nodes with regular traffic between
neighbors and a “front” of the chain at any point in time.
Onward colonization at the front would have been achieved
by individuals or groups seeking out new habitat and/or being
blown along the coast. The capacity would always have been
present for community members to explore unfamiliar coast-
line until they could return with news of a suitable estuary or
until their water dwindled to a bare sufficiency for the return
journey—a version writ small of Irwin’s (1992) model for the
colonization of the Pacific. Thus, expansion of the social milieu
at its front would have been a function of maintaining the
social milieu, equipped with technology that both held society
together and allowed it to expand. By assuming serviceable
watercraft, portage of potable water, and an estuarine focus, no
further assumptions would be necessary to underwrite the
rapid-southern-dispersal hypothesis.
Watercraft and Estuaries in the Southern Dispersal
We may assume that early watercraft was paddle-propelled,
limiting its role in rapid transport to rare or inadvertent oc-
casions. Experienced canoe paddlers today can achieve merely
6–18 miles (10–30 kilometers) per day, depending on wind
conditions (Search and Rescue Society of British Columbia
2006). This suggests a maximum allowance of 10 kilometers
on a usual day for Late Pleistocene paddlers, depending on
the quality of their craft—in any case, a shorter distance than
could be achieved by walking. However, strong winds and
especially gales enforce downwind travel on any paddle craft
caught in the open. Regular users of coastal watercraft would
have needed to be prepared for occasions when all they could
do was hold tight during a rapid maritime journey.
One major advantage boat users have over pedestrians is
the much lower friction of water compared with land. A boat
can double as a mobile shack without incurring prohibitive
transport costs. The vagaries of the wind would have exerted
selection pressures on boat users to carry containers for fresh
water along with gear for foraging and tool-kit maintenance.
These safeguards would additionally have allowed those near
the dispersal front to skirt shores bereft of even the low-tide
water seeps mentioned by Sauer (1963). Stretches of 100 ki-
lometers or more of the most forbidding coast could have
been traversed by single colonists or small cooperative groups,
intentionally or otherwise. The most critical factor for the
harshest coasts would have been to start the journey with
decent water supplies. This point suggests that we should look
at the estuary, defined by Nybakken (2001, 328) as “a partially
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Current Anthropology Volume 48, Number 2, April 2007
Usefulness of a Darwinian Perspective
coastal exploitation with simple watercraft would have been
greatest at times of rising sea levels, when reefs, mangrove
belts, and estuaries would have been richest and in closest
proximity to each other. They accordingly indicate
59,000–62,000 years ago as the optimum time for colonization
of Australia across the Timor Sea. This dating is in complete
accord with the rapid-southern-dispersal hypothesis, and the
newcomers’ tradition of watercraft and an estuarine focus, as
suggested here, would have preadapted them for the rigors
of the southern route. 1
The advantages of the northern route include intervisibility
between islands all the way to New Guinea, superior fresh
water provisioning, and an eastward direction of the pre-
vailing winds (Mulvaney and Kamminga 1999, 108). However,
the profusion of debouching streams along the coasts of
Sunda and the wetter Wallacea islands would have blurred
the distinction between estuarine and other habitats. When
human groups with a narrow niche breadth (as posited here)
came upon a wider range of options, how would they have
responded? The most likely answer is variably. Some groups
may have used the safety net of ready drinking water to un-
derwrite even more ambitious exploration of the surrounding
coasts and seas. These would have been the first colonists of
the well-watered coasts, explaining the “maritime nomad”
adaptation (Gosden and Robertson 1991; see also Torrence
et al. 2004) that emerges in the earliest archaeology of the
Bismarck Archipelago. Other groups, however, would have
settled for the estuaries at hand, filling in the coastline with
directionally irregular movements. Of course, filling in coast-
lines, river fronts, and the landscape generally would have
been the major solution to population growth anywhere be-
hind the colonizing front, albeit involving the slower process
of individual learning.
Pardoe (2006) suggests that Sahul could have been colo-
nized through one or more organized expeditions, perhaps
numbering thousands of transmigrants. This is possible but
less parsimonious than a scenario of disorganized coloniza-
tion. The considerable number of Wallacea islands within
striking distance of Sahul (fig. 1) suggests multiple sources
and destinations. Particularly with the likelihood of periodic
genetic replenishment, even a small band of arrivals may have
thrived much better than would be suggested by Moore’s
(2001) population genetic models. This is because the incest
prohibitions assumed by Moore relate specifically to filled
landscapes, not to isolated bands short of unrelated marriage
partners but very well endowed with land. There would have
been a dynamic balance between the enhanced ability of rel-
atively small parties to make early landfall and the greater
viability of larger populations of colonists (Webb 2006,
98–101). The original contribution which the present model
A Darwinian perspective would remove any need to invoke
human teleological intentionality either to exploit the synergy
between watercraft and estuarine habitation or to plan their
combination for purposeful colonization. A Darwinian par-
adigm characterizes cultures by their stream of experimental
innovation within the social and technological limits of pre-
vailing social practices (Rindos 1986). As is the case with
genetic mutations, random processes strongly affect the in-
ception and establishment (“fixation”) of innovations, but if
they survive the initial trials their frequency will increase or
decrease under the natural selection mechanisms that operate
on cultural traits (Boyd and Richerson 1985, 2005). In par-
ticular, it is not necessary for an estuarine focus to have stim-
ulated innovation in watercraft or vice versa. As long as a
tradition combining watercraft and estuarine habitation hap-
pened to develop in some population, the adherents would
have reaped the benefits. The incubation period may have
been selectively neutral or even maladaptive from a func-
tionalist point of view; Boyd and Richerson’s (1985, chap. 8)
model of “runaway cultural evolution” explains how cultural
evolution can, within limits, take on its own life independently
of the reproductive success of the culture bearers.
East Africa, with its record of littoral and reef exploitation
by 125,000 years ago, was a likely nursery or welcome recipient
of the watercraft-estuarine tradition at some point during the
succeeding 50,000 or so years. A “rapid” dispersal of this
tradition along the northern rim of the Indian Ocean would
have been aided by the similarity of coastal resources along
the entire route. Hinterland colonists, in contrast, would have
been severely slowed down by the need for “individual learn-
ing,” the slow process whereby Darwinian selection leads to
new modes of successful behavior in a novel environment
(Boyd and Richerson 2005, pt. 1).
Colonization of Sahul
The colonization of Sahul could have occurred along a north-
ern route through Sulawesi or a southern route via Nusa
Tenggara (fig. 1). Whichever route was followed, colonization
had presumably occurred in the search for habitable islands
rather than as a heroic thrust from Sunda to Sahul. Travel
between islands could have been accelerated by paddling to-
wards parts of the sea with tell-tale signs of land nearby or
by being swept away by tides and storms (Mulvaney and
Kamminga 1999, 107–12). With regard to the model of es-
tuarine seeking developed here, preparedness for short voy-
ages over open sea would need to be added to the coastal
travel that had brought Homo sapiens to the edge of Wallacea.
An estuarine focus would have been particularly beneficial,
if not obligatory, for movement along the typically dry shores
of Nusa Tenggara (Monk, de Fretes, and Reksodiharjo-Lilley
1997, 76–77) and northwestern Australia. O Connor and
Chappell (2003, 21) point out that the opportunities for
1. The previous period of higher sea levels, according to O’Connor
and Chappell (2003, fig. 2), would have prevailed from
70,000
skirted Arabia according to the rapid-southern-dispersal hypothesis.
∼ 75,000 ∼
years ago, just when Homo sapiens departed East Africa and
to
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319
may make to understanding the colonization of Sahul is to
suggest a possible motive for exploration—seeking out new
estuaries—and to assign colonists a habitat search image that
would have optimized their chances of survival upon landfall.
model that did not specifically rule out use of coastal re-
sources. Conceivably, after recovering a satisfactory sample of
uplifted Indian Ocean coastal sites dated to 60,000–75,000
years ago, archaeologists may be able to determine a pattern
of increasing or decreasing site density towards ancient es-
tuaries, either consistent with or falsifying the model. This
would be a tall order but perhaps not impossible given on-
going advances in geoarchaeology and dating techniques.
Three current anomalies in Australasia’s archaeological rec-
ord can perhaps be explained by an estuarine focus. First,
O’Connor and Chappell (2003) point out that the terminal
Pleistocene in Australia seems to have witnessed colonization
from savanna to coast rather than the reverse. They do not
doubt that the initial colonists had a maritime capacity, but
they see no evidence that this had confined early dispersal to
littoral belts. My model would provide an archaeologically
undetected means for colonists to have discovered and settled
estuaries, moved upriver, and adapted to savanna resources
(cf. Bowdler 1990). Secondly, early H. sapiens colonists may
have had limited advantages over earlier established Eurasian
populations except at estuaries (or perhaps the coastline gen-
erally) which they commandeered. Modeling early H. sapiens
more as supertramp (cf. Diamond 1977) than as superman
could explain the occupation of the Flores interior by H.
floresiensis till the end of the Pleistocene (Morwood et al.
2005; Brumm et al. 2006). Thirdly, despite passage through
Wallacea before 50,000–60,000 years ago, a true maritime
culture evidently appeared there only after 30,000–35,000
years ago (Bellwood 1998). The explanation provided by the
present model is that the colonists had more an estuarine
than a maritime orientation. The possibility of 50,000-year-
old sites in Sulawesi, along one of its main rivers (Keates and
Bartstra 1994), is particularly interesting in this regard.
Effective testing of the rapid-southern-dispersal hypothesis
can be expected from ongoing genetic research and its cor-
relation with growing archaeological knowledge. Merriwether
et al. (2005) note that their evidence on M haplogroups may
be consistent with the rapid-dispersal hypothesis of Macaulay
et al. (2005 a ), but it could also be explained by the existence
of ancestral hapolotype M for around 20,000 years. Should
further research bear out the latter explanation, the rapid-
southern-dispersal hypothesis would require revision. In ad-
dition, as previously implied, expert opinion is currently split
between
Testing of the Model and Its Implications
for Archaeology
The present model posits a process of colonization of estu-
arine habitats between approximately 75,000 and 60,000 years
ago. Any sites dating to the last 40,000 or 50,000 years ago,
when radiocarbon dating comes in, would refer to adaptations
postdating the colonization thrust by 10,000 to 35,000 years.
The relevance of these sites to the testing of the model would
be debatable. It may draw support from radiocarbon dates
of around 40,000 years ago at Kupona na Dari in New Britain
(Torrence et al. 2004) and Buang Merabak in New Ireland
(Leavesley et al. 2002), but alternative colonist models would
be equally comfortable with the same archaeological evidence.
Absence of evidence of 75,000–60,000-year-old sites along
the northern rim of the Indian Ocean would hardly amount
to evidence of their absence. Late Pleistocene coastal sites are
rarely available for archaeological study except where the land
has risen faster than the sea, as with the Bismarck sites, or
where a steep continental shelf translates into a short distance
from the Pleistocene coast (e.g., Mandu-Mandu in Western
Australia). Other coastal sites would have been destroyed
through wave action or, at best, buried beneath marine sed-
iment. While we may not have a single site in Arabia or
southern Asia that documents use of the coast between 60,000
and 75,000 years ago, South Asia has possible candidates,
including Badalpur (Marathe 1981, 47, 111–12; see James and
Petraglia 2005, S7), Patirajewalla Site 50 (Deraniyagala 1992,
685; Abeyratne 1996, 101), and Ramayogi Agraharam (Rath,
Reddy, and Prakash 1997). This small number should be seen
in the context of the general rarity of South and Southeast
Asian archaeological sites dated to around 60,000 years ago
(Bellwood 2001; James and Petraglia 2005; Mellars 2006). To
falsify the southern-dispersal hypothesis, we would need pos-
itive evidence of avoidance of the relevant stretch of coast.
This could come from archaeologically sterile deposits, se-
curely dated to the period concerned, in a location where we
could confidently expect traces of any coastal habitation. I
am unaware of any documented case to this effect.
The prospects for archaeological testing are further dimmed
when we consider the estuarine focus of the model. During
times of falling sea levels such as the Late Glacial Maximum,
rivers would have cut down to lower levels, eroding sites along
their banks or near their mouths. Moreover, this destruction
of estuarine sites would have been most pronounced where
tectonic uplift would otherwise have lent these sites some
chance of survival. To be sure, an estuarine focus hardly im-
plies avoidance of the adjacent coast, so the model would
predict sites on uplifted land adjacent to ancient estuaries.
However, so would any other habitation or colonization
50,000
Australia. Should the former estimate become the consensus
and the departure date from East Africa of
and 60,000 years ago for initial entry to
75,000
ago still hold up, this would imply a less than rapid dispersal
to Sunda or a stasis at Sunda, either of which would be in-
consistent with the model developed here.
years
Conclusion
This contribution develops an estuarine colonist model to
address and explain the rakelike structure currently observed
for the most deeply rooted mtDNA lineages outside of Africa.
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