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Quaternary International 300 (2013) 32e47
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Quaternary International
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Human dispersal across diverse environments of Asia during the
Upper Pleistocene
Nicole Boivin a, Dorian Q. Fuller b, Robin Dennell c, Robin Allaby d, Michael D. Petraglia a, *
a
School of Archaeology, Research Laboratory for Archaeology and the History of Art, University of Oxford, Dyson Perrins Building, South Parks Road,
Oxford OX1 2HU, UK
b
Institute of Archaeology, University College London, 31-34 Gordon Square, London WC1H 0PY, UK
c
Department of Archaeology, University of Sheffield, Northgate House, West Street, Sheffield S1 4ET, UK
d
School of Life Sciences, University of Warwick, Wellesbourne, Warwick CV35 9EF, UK
a r t i c l e i n f o
a b s t r a c t
Article history:
Available online 15 January 2013
The initial out of Africa dispersal of Homo sapiens, which saw anatomically modern humans reach the
Levant in Marine Isotope Stage 5, is generally regarded as a ‘failed dispersal’. Fossil, archaeological and
genetic findings are seen to converge around a consensus view that a single population of H. sapiens
exited Africa sometime around 60 thousand years ago (ka), and rapidly reached Australia by following a
coastal dispersal corridor. We challenge the notion that current evidence supports this straightforward
model. We argue that the fossil and archaeological records are too incomplete, the coastal route too
problematic, and recent genomic evidence too incompatible for researchers not to remain fully open to
other hypotheses. We specifically explore the possibility of a sustained exit by anatomically modern
humans, drawing in particular upon palaeoenvironmental data across southern Asia to demonstrate its
feasibility. Current archaeological, genetic and fossil data are not incompatible with the model presented,
and appear to increasingly favour a more complex out of Africa scenario involving multiple exits, varying
terrestrial routes, a sub-divided African source population, slower progress to Australia, and a degree of
interbreeding with archaic varieties of Homo.
Ó 2013 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
Archaeological discoveries in the Levant have long played a
critical role in debates about human evolution. Situated at the
western edge of the Asian landmass, the small region has, since the
late 1920s, yielded key fossil finds of both Homo sapiens and Homo
neanderthalensis (Shea, 2003, 2008). Interpretation of the Levantine
sites has never been straightforward, however, with problems of
chronology and a shared Mousterian technology, for example,
generating numerous opportunities for controversy. While findings
in the region were initially taken to support a gradual transition
between H. neanderthalensis and H. sapiens, improved chronologies
have indicated significant overlap between the two types of human,
on the order of 25e50,000 years. This recognition has seen the
Levantine fossil remains come to be drawn upon as a major strand
of evidence in support of a replacement model for modern human
origins, with the Levantine H. sapiens fossils now widely interpreted as representing a first foray of modern humans out of Africa,
* Corresponding author.
E-mail address: [email protected] (M.D. Petraglia).
1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved.
http://dx.doi.org/10.1016/j.quaint.2013.01.008
at a time of improved climatic conditions in Marine Isotope Stage 5
(MIS 5) (Frumkin et al., 2011). In the absence of early fossil finds
further afield and in light of certain primitive morphological features, the Levantine MIS 5 exit has generally been seen as a failed
dispersal (Mellars, 2006a, 2006b; Shea, 2008; Oppenheimer, 2009,
2012). This notion is offered support by recent molecular analyses
of mitochondrial DNA (mtDNA), which suggest a successful
expansion of H. sapiens out of Africa only at around 60 thousand
years ago (ka) (e.g. Macaulay et al., 2005; see below).
The apparent agreement of archaeological and genetic findings
has led to the emergence of a consensus view regarding the dispersal of modern humans out of Africa. This view holds that
H. sapiens left Africa in MIS 4 or early MIS 3, and colonized Australia
within perhaps as little as several thousand years by rapidly skirting the coasts of the Indian Ocean. While this may be a reasonable
hypothesis, its uncritical and increasingly widespread multidisciplinary acceptance suggests the need for more careful
appraisal. Here we will first examine current fossil, archaeological
and genetic findings concerning the dispersal of H. sapiens from
Africa, critically appraising these sources of evidence. We then
consider how this evidence sits in relation to palaeoenvironmental
evidence and a hypothesized dispersal of modern humans across
N. Boivin et al. / Quaternary International 300 (2013) 32e47
southern Asia. As we discuss below, a number of sites in the Arabian
Peninsula and South Asia have begun to yield Middle Palaeolithic
assemblages dating to MIS 5, suggesting that H. sapiens may have
moved into these geographic zones during climatic ameliorations.
The archaeological and palaeoenvironmental evidence is supported
by new genetic findings that challenge the orthodox view that
H. sapiens replaced archaic species of Homo in Asia. While there
seems little need to invoke a multiregional model for the evolution
of H. sapiens to explain these new genetic findings, more nuanced
models of population structure and a degree of assimilation are
required to explain a suite of recent autosomal and ancient DNA
studies (to which we return to below). Overall, the picture of
modern human origins, which seemed at one point to be crystallising into a fairly comfortable orthodoxy, is now looking decidedly
more complex (Dennell and Petraglia, 2012). The time is ripe for
consideration of alternatives to a straightforward MIS 4 or 3 exit of,
and complete replacement by, modern humans, and in particular to
examine the plausibility of earlier exits across southern Asia during
MIS 5.
2. Absence of fossils and artefacts
Evidence for the late dispersal of modern humans out of Africa,
and replacement of local archaic species of Homo, is by no means
straightforward to interpret. Outside the sphere of molecular
genetics, the story rests entirely on evidence from fossils and artefacts. Each class of evidence has its own problems, but both suffer in
particular from regional biases in archaeological research. While
industrialised regions with long histories of archaeological research
like Europe and the Levant have seen more comprehensive study,
equally if not more critical regions like the Arabian Peninsula, South
Asia, East Asia and Southeast Asia have received relatively uneven
coverage, and in some cases, minimal investigation.
The earliest well-dated fossils of modern humans outside of
Africa, excluding the Levant, are in Laos (c. 46 ka) (Demeter et al.,
2012), Sunda (c. 46 ka) (Barker et al., 2007), Australia (c. 40 ka)
(Bowler et al., 2003), Sri Lanka (c. 38 ka) (Deraniyagala, 1992; Perera
et al., 2011) and India (c. 20 ka) (Kennedy, 2000). In Arabia, modern
human fossils have not been dated to earlier than the Holocene
(Martin, 2007). Taken at face value, these dates might be interpreted to indicate a dispersal of modern humans from east to west
rather than from out of Africa. The paucity of well dated fossils has
not, however, prevented palaeoanthropologists from drawing
meaning from the observed patterns. The earliest dated fossils from
Sunda and Sahul are frequently taken to suggest not a minimal date
for the exit of modern humans out of Africa, but an approximate
date for Out of Africa. Since all the other dates for intervening
regions in the west are younger, and thus clearly unreliable as
indicators of initial dispersal, the dates from Sunda and Australia, in
particular, have come to assume significance in attaching a chronology to the narrative of Out of Africa. It needs to be emphasised,
however, that the earliest fossils do not necessarily date Asia’s first
modern human colonization, let alone reflect a date for the exit of
H. sapiens from Africa, contrary to some influential models that
have been proposed.
Artefact assemblages also present their own challenges.
Archaeological sites are subject to alterations and transformations
in depositional contexts, and these play out exceptionally harshly in
the hyper-arid and tropical environments that today dominate
some of the most plausible out of Africa dispersal routes. Even the
most well-preserved objects, stone tools, are often difficult to find
in well stratified, datable contexts e the Arabian Peninsula, for
example, has yielded a wealth of Palaeolithic surface finds
(Petraglia and Alsharekh, 2003), but until recently, very few
stratified assemblages (Groucutt and Petraglia, 2012).
33
Most pressing of all, however, is the problem of identifying
straightforward signatures of modern human behaviour. In
accordance with traditional European-derived models, the arrival
of modern humans in a particular region was once seen to be traced
relatively easily, through the appearance in the archaeological
record arrival of evidence for art, symbolism, and complex Upper
Palaeolithic technologies, as seen in Europe around 45e40 ka.
Recent years have, however, seen a thorough deconstruction of
the notion of a straightforward ‘modern human revolution’. It has
been recognised that so-called modern human behaviours, as
traced through their apparent archaeological manifestations: 1) do
not necessarily arrive as part of a package; 2) do not necessarily
exhibit continuity through time; and, most significantly, 3) are not
always present in modern human populations. There is nothing in
terms of material culture attributes that is unique to H. sapiens and
also universal among modern humans in Africa, Eurasia and Australasia. It is thus the case that with material culture as with fossils,
absence of evidence need not necessarily indicate evidence of
absence.
Contrary to the notion of the sudden revolutionary appearance
of a new behavioural package with the emergence of modern
humans, a major synthesis of behavioural indices for Middle Stone
Age Africa revealed the gradual and non-contiguous emergence of
modern traits (McBrearty and Brooks, 2000). Other parallel syntheses for South Asia (James and Petraglia, 2005) and Sahul
(Brumm and Moore, 2005; O’Connell and Allen, 2007; Habgood and
Franklin, 2008) have revealed a similar trend. The Australian evidence is particularly interesting in that H. sapiens appears to have
arrived without an African ‘package’ of innovations, and to have
acquired these independently and piecemeal. Examination of the
South-East Asian record has also demonstrated the inadequacy of
traditional markers of modern human behaviour, such as refined
blade and bladelet technology, body ornamentation and mobiliary
and parietal art (Barker et al., 2007; Moore and Brumm, 2007), and
has focused instead on the appearance of, for example, new subsistence strategies and modes of engagement with the landscape
with the arrival of H. sapiens (Summerhayes et al., 2010). Such
studies have demonstrated the Eurocentric biases of models of
behavioural modernity, which created a set of global expectations
from a regional record. In reality, there is no single material trait
that is inevitable in modern human assemblages.
What this means for Out of Africa models is that cognitive
modernity may have emerged simultaneously or even before
(Kingdon, 1993; Foley and Lahr, 1997) morphological modernity,
and the appearance of traditional indicators of behavioural modernity in the archaeological record may mark demographic or social
changes (Shennan, 2001; Petraglia et al., 2009; Powell et al., 2009),
rather than cognitive ones. These new ideas about modern human
behaviour suggest the possibility that H. sapiens may have been
fully modern back to 200 ka or more, and exited Africa prior to the
appearance of, or without bringing along, more complex technologies or archaeologically-attested symbolic practices. We cannot therefore rely on these forms of material evidence to trace
dispersals any more than we can rely on fossils as indicators of the
first anatomically modern humans to leave Africa. New understandings of behavioural modernity furthermore open up the
possibility that the H. sapiens populations that exited Africa and are
documented in the Levant in MIS 5 may have been fully cognitively
and behaviourally modern (contra Mellars, 2006b).
3. Coastal route out of Africa?
Another problem with the current consensus view for Out of
Africa is its increasing focus on a coastal route of dispersal. Part of
the problem is that two related but separate hypotheses have
34
N. Boivin et al. / Quaternary International 300 (2013) 32e47
frequently been conflated. One argues that, along with a northern
exit towards Europe and Siberia around 45 ka, H. sapiens also followed a more southerly route out of Africa at an earlier date, dispersing through southern Asia (Lahr and Foley, 1994, 1998). The
other suggests that modern humans ‘coasted’ out of Africa
(Stringer, 2000; Bulbeck, 2007), travelling along coastal corridors
around the Indian Ocean rim to reach Australia by 45 ka. The latter
idea has a long history, appearing in various manifestations since
Sauer (1962) suggested the potential role of coastlines as corridors
of dispersal for early humans. Subsequent discussions by Kingdon
(1993), and by Field and colleagues (Field and Lahr, 2005; Field
et al., 2007), explored the possibilities of a southern, coastallyfocused dispersal route for modern humans out of Africa, all or
part of the way along the Indian Ocean rim. The idea of a coastal
route has been embraced by a number of geneticists, who have
drawn upon it to explain certain patterns in the genetic record of
human diversity (Macaulay et al., 2005; Thangaraj et al., 2005;
Oppenheimer, 2009).
The gist of the coastal dispersal hypothesis (which should be
distinguished from the more generic southern dispersal hypothesis) is that maritime environments offered particularly favourable
environments for human occupation, and provided a dispersal
corridor that enabled rapid movement along the Indian Ocean rim
of initial colonising populations. This single dispersal event corresponded with the arid conditions of MIS 4 (Lahr and Foley, 1994;
Stringer, 2000; Forster and Matsumura, 2005; Mellars, 2006a;
Bulbeck, 2007; Oppenheimer, 2009), when desert dominated in
northern Africa, the Arabian Peninsula, and the northwestern part
of the Indian subcontinent. Such a model argues that humans were
able to draw upon the rich resources of coastal environments,
manifesting a maritime adaptation that led them to rapidly skirt
continents (Oppenheimer, 2009), and to develop the seafaring
technology that eventually enabled colonisation of Sahul (Forster
and Matsumura, 2005). Research on the Red Sea coast of Eritrea
and in coastal South Africa has suggested human adaptation to a
coastal marine environment by MIS 5, and indicated to some that
the emergence of modern human behaviour may be linked to a
more intense and complex exploitation of marine resources
(Deacon, 1989; Walter et al., 2000; Marean, 2010). Several recent
genetic studies of mtDNA diversity, meanwhile, have suggested a
single exit of modern humans out of Africa, as well as a rapid dispersal to Australia (Forster, 2004; Forster and Matsumura, 2005;
Macaulay et al., 2005; Oppenheimer, 2009). These features have
been seen as consonant with a coastal route, both because coasts
are increasingly viewed as a kind of ‘highway’, enabling rapid dispersal, but also because desert expansion in many inland regions
during the relevant time period would have created barriers to
dispersal, forcing populations to move along coasts.
Despite the popularity of the coastal dispersal hypothesis, there
are numerous reasons why the idea should be approached with a
greater degree of caution. One is that archaeological evidence
indicative of a maritime adaptation in early humans is extremely
limited. At present, relatively secure evidence for use of marine
resources by early H. sapiens appears in South Africa (Marean,
2010). Shell beads found in North Africa (d’Errico et al., 2009)
indicate visitation to the coast and not coastal adaptation necessarily. The claim for an association between marine resources and
archaeological sites in Eritrea (Walter et al., 2000) also include lag
deposits and natural death assemblages (Bruggemann et al., 2004),
which do not necessarily demonstrate human subsistence activities
(Bailey, 2009). Even if we accept that a maritime adaptation is
plausible, however, problems remain with the model. Bailey (2009)
has noted the implausibility of the idea that the emergence of
modern human cognition ushered in a sudden awareness of the
richness of coastal resources, whose consumption was
subsequently of an intensity sufficient to lead to local overexploitation and to ultimately spur dispersal. Not only is exploitation of marine resources sufficiently simple to have been
competently undertaken by archaic humans (Cortés-Sánchez et al.,
2011) and even non-human primates (Malaivijitnond et al., 2007),
the probability that humans were at this early stage marine specialists is, based on ethnographic and archaeological parallels,
extremely low (Bailey, 2009). The many inland Upper Pleistocene
sites in Arabia, South Asia, East Asia and Southeast Asia argue much
more for a more flexible subsistence approach in which terrestrial
plant and animal resources continued to play a major role in human
lifestyles. Even early modern human sites that are described as
‘coastal’ may furthermore demonstrate a terrestrial focus. Despite
being one of the Upper Pleistocene human occupation sites closest
to the coast, for example, Niah Cave in Borneo possesses a faunal
assemblage dominated by terrestrial and freshwater taxa, and lacks
evidence for the use of marine resources (Barker et al., 2007).
Coastal sites may indeed be lacking from the record due to submergence along shallow continental shelves in many areas,
although surveys in areas with tectonically raised reef formations
and former shorelines have not produced evidence for marineoriented archaeological sites either (Korisettar, 2007; Bailey,
2009). Even if we grant that most sites are submerged, the presence of many inland sites stresses the importance of terrestrial
foraging adaptations over the marine specialization that is implicit
in coastal dispersal models, particularly those that imply a rapid
dispersal where repeated overexploitation of resources led to
movements further down the coast.
Equally problematic is the idea that coasts were particularly rich
and attractive environments for early humans. While certain
coastal environments are favourable habitats, other coastal areas
are effectively ‘deserts’ lacking entirely in critical freshwater
resources. The notion of a ‘coastal oasis’ e the idea that freshwater
springs appeared on emerged continental shelves at times of low
sea level (Faure et al., 2002) e is central, often implicitly, to most
models of coastal dispersal, yet lacks empirical support (Bailey,
2009). Even if valid, the coastal oasis model does not postulate
the kind of uniformly equable coastline that such a dispersal
necessitates (Sauer, 1962; Stringer, 2000). Instead, it suggests the
probability of rather dispersed and non-interconnected wetland
environments in places where underground springs exist. There are
also many coastal regions wholly devoid of shellfish or any obvious
marine sources that could be easily gathered or otherwise obtained.
Furthermore, the idea that coasts were, in contrast to inland areas,
particularly stable environments is open to question. Westley and
Dix (2006) summarise evidence for coastal instability during the
Upper Pleistocene, and emphasise that present day coastlines, and
our ideas about coasts, are the product of relatively unique environmental circumstances during the exceptionally stable Holocene
interglacial.
In sum, while coastal regions may have been used as dispersal
routes in prehistory, both for H. sapiens and earlier hominid exits
out of Africa, the notion of a coastal superhighway between Africa
and Australia that enabled unexpectedly fast dispersal is simplistic
and derives little if any support from empirical data. While probably a key necessary corollary of the currently popular notions of an
MIS 4 exit for Out of Africa, and a rapid dispersal to Australia, it may
be argued that the coastal dispersal model has a number of significant deficits and unproven assumptions.
4. The genetics of Out of Africa
If fossil and archaeological evidence offer little support for a
rapid coastal dispersal of H. sapiens out of Africa at c. 60 ka, genetic
data is equally problematic. While a consensus date of c. 60 ka is
N. Boivin et al. / Quaternary International 300 (2013) 32e47
often cited for the mtDNA data, recent literature actually offers a
range of dates for human expansion out of Africa, anywhere
between 85 and 45 ka (Oppenheimer, 2003, 2009, 2012; Macaulay
et al., 2005; Kivisild et al., 2006; Cabrera et al., 2009; Endicott et al.,
2009; Soares et al., 2009; Rasmussen et al., 2011). Variation is due,
in part, to the values used to calibrate the mtDNA clock (e.g., Scally
and Durbin, 2012). The problems associated with calibrating the
mtDNA clock have been noted, including in particular the reliance
on problematic assumptions about the timing of the divergence
between humans and chimpanzees, as well as the constancy of the
molecular clock (Ho and Larson, 2006; Endicott et al., 2009). Further research is required to improve confidence in molecular estimates of human evolutionary timescales.
Another under-appreciated issue is the anomalous nature of the
genetic evidence for a rapid spread of modern humans from Africa
to Asia. Echoing the fossil date anomaly, the mtDNA branch lengths
for sampled populations are longest for those which are farthest
east, in Near Oceania, and shortest in the Asian areas that
would have been encountered first (Merriwether et al., 2005;
Oppenheimer, 2009). The real problem, however, is that the variation in branch lengths suggests that a single genotype engaged in
the expansion actually existed for 30 ka, which does not support a
rapid expansion. The anomaly can be explained by what we call an
‘M buffer’ effect (see Supplementary material A) which implies that
the branch ages we observe are considerable underestimates of the
time of arrival of the genotype to these areas. Such anomalously
long-lived genotypes have been directly observed through ancient
DNA in species such as the Iberian lynx (Rodríguez et al., 2011).
Much more problematic, however, is the challenge posed by
new genomic studies that offer perspectives on evolutionary history that sometimes contrast markedly with those of mitochondrial
and Y-chromosome-based investigations. Recent analysis of the
frequency of alleles of SNPs (single nucleotide polymorphisms)
from the International HAPMAP Project, for example, suggests that
the out of Africa expansion may actually need to be modeled as two
or more bottlenecks, rather than the single bottleneck suggested by
mtDNA studies (Keinan et al., 2007; Wall et al., 2009). The authors
of one study (Keinan et al., 2007) suggest that a two bottleneck
model provides the potential best fit to the SNP data, with the first
at w140e80 ka, potentially reflecting the MIS 5 exit identified in
the Levant, and the second dating to the Last Glacial Maximum and
potentially associated with demographic upheavals at this time. A
bottleneck dating to anywhere around 60 ka is not presently supported by such studies.
Even more challenging is the presence of autosomal genes
whose estimated ancestry extends well beyond the conventionally
accepted date for the speciation of modern humans. While a single
origin model predicts that all shared polymorphisms trace their
origins back to a single deme in Africa, several regions of the
nuclear genome, both on the X chromosome and on the autosomes,
have deep ancestries that are incompatible with a simple, single
origin model (Garrigan and Hammer, 2006). These include, for
example, the RRM2P4 region of the X chromosome that has an
estimated Time of the Most Recent Common Ancestor (TMRCA) of
2.33 million years and an apparent East Asian ancestry (Cox et al.,
2008) and the microcephalin D allele, which has a TMRCA of 1.1
million years (Evans et al., 2006). These kinds of findings do not
support a straightforward single origin model (see also Harding
et al., 1997; Eswaran et al., 2005; Harpending and Eswaran, 2005;
Relethford, 2008; Weaver and Roseman, 2008), and have been
interpreted as reflecting population structure within Africa and/or
interbreeding with archaic species of Homo outside of Africa. Certain mitochondrial (Behar et al., 2008) and autosomal (Harris and
Hey, 1999; Labuda et al., 2000; Harding and McVean, 2004; Satta
and Takahata, 2004; Barreiro et al., 2005; Garrigan et al., 2005;
35
Plagnol and Wall, 2006) (but see Li et al., 2008) studies suggest that
populations in Africa were structured, with numerous small and
separately evolving populations present in the continent at the
time of the out of Africa dispersal. Metapopulation models, which
argue for the presence of multiple populations, and their absorption of some admixture before neighbouring extinctions, are more
compatible with the wide range of TMRCA estimates for autosomal
loci than the straightforward single origin model of out of Africa
(Harding and McVean, 2004). Cranial data offer support for a more
complex out of Africa scenario involving multiple dispersals,
probably by varying routes, at different times and/or from a subdivided African population (Lahr, 1996; Schillaci, 2008; Gunz et al.,
2009).
Archaic admixture in the modern human genome in Eurasia is
now clear. While studies of mtDNA (Richards et al., 1996; Krings
et al., 1997; Handt et al., 1998; Serre et al., 2004; Green et al.,
2008) and the Y-chromosome (Jobling and Tyler-Smith, 2003)
have not indicated admixture, for example, it was widely recognized that autosomal DNA might tell a different story. Indeed this
has proven to be the case, with evidence of archaic ancestry surfacing for the microcephalin gene (Evans et al., 2006), the tau
(MAPT) locus (Hardy et al., 2005), the b-globin gene (Harding et al.,
1997), the immunity genes CD209 and CD209L (Barreiro et al.,
2005), the dystrophin gene (Labuda et al., 2000) and others (Wall
et al., 2009). Direct contact between archaic and modern humans
is also supported by autosomal data for modern human head lice,
Pediculus humanus (Reed et al., 2004). More recently, a draft
sequence of the Neanderthal genome has been published that
indicates gene flow from Neanderthals into the ancestors of nonAfricans (Green et al., 2010). The study also demonstrated that
Neanderthals shared more genetic variants with present-day
humans in Eurasia than with present-day humans in sub-Saharan
Africa, suggesting that the gene flow occurred before the divergence of Eurasian groups from one another. The main known region
in which H. sapiens and H. neanderthalensis co-existed prior to the
separation of European and Asian lineages of human is the Levant
(but see Shea, 2008). Given that the overlap between these species
in the Levant may have lasted only until c. 75 ka, this might be seen
as supporting an exit in MIS 5. However, as our distribution map for
Neanderthals (Fig. 1; Supplementary material B) suggests, a broad
swath of western and central Eurasia may have been home to
Neanderthals from MIS 5 until MIS 3, suggesting a range of possible
regions of co-occupation. Subsequent regionally specific admixture
is suggested by the presence of ancient alleles that are found only or
predominantly in certain regions e for example, certain Europeanlocalised alleles for microcephalin in Europe (Evans et al., 2006),
and Asia-localised alleles for b-globin dating to >200 ka (Harding
et al., 1997) e and by the presence of basal clades composed
entirely of Asian sequences, as for the RRM2P4 region (Cox et al.,
2008). The Neanderthal findings were soon after accompanied by
a second hominin draft genome, that of the Denisovans of Siberia
(Krause et al., 2010; Meyer et al., 2012). The finding of gene flow
from Denisovans into some modern human populations of Melanesia suggest that admixture occurred after the dispersal of
H. sapiens into Asia, and probably somewhere in South or Southeast
Asia (Stewart and Stringer, 2012).
Recent revision of the human genomic mutation rate provides
another line of nuclear DNA evidence that problematizes mtDNAbased Out of Africa estimates, while at the same time offering a
chronology that supports evidence for modern human and archaic
admixture. Direct measurements of the nuclear genomic mutation
rate in contemporary humans using next generation sequencing
technology have indicated a value that is approximately half of that
previously estimated from fossil calibration (Scally and Durbin,
2012). When applied to the analysis of the time of separation of
36
N. Boivin et al. / Quaternary International 300 (2013) 32e47
Fig. 1. Hominin populations during MIS 5. H. sapiens was probably the main hominin resident in Africa by this time. Neanderthals prior to MIS 5 were distributed in Europe and the
Levant, but expanded their range across Asia thereafter (Howell, 1999). H. erectus sensu stricto was probably confined to East Asia by MIS 5 as the Ngandong H. erectus assemblage
now appears to have been older (Indriati et al., 2011). Because of the lack of diagnostic skeletal evidence from Arabia and South Asia between 125 and 40 ka, the identity of their
inhabitants is unknown, but, we argue, likely to have been H. sapiens. The identification of H. sapiens in Laos helps to support this argument. Note the possible areas of overlap
between H. sapiens and the southern limits of Neanderthals and H. erectus which would allow for interbreeding e as indicated by recent studies and possibly by fossil evidence from
Zhirendong and other Chinese caves. The distribution of “Denisovans” is unknown as they are indicated only by the ancient DNA of a phalange from Denisova Cave, Siberia, and
modern genetic studies of SE Asians and Melanesians. Present indications are that they are probably an East Asian sister population of Neanderthals, but little more is known at
present. For Neanderthals, the purple area denotes their likely “core” distribution. The adjoining blue areas show their likely maximal extent, northwards in warmer episodes, and
southwards when conditions in the core area deteriorated, or when opportunities arose for expansion. Their easternmost distribution towards NE China and the Pacific coast of NE
Asia is shown as uncertain. Filled circles: H. sapiens: 1) Jebel Irhoud, Morocco; 2) Skuhl, Qafzeh, Israel; 3) Herto, Ethiopia; 4) Niah, Borneo; Filled stars: Neanderthals: 5) Tabun,
Kebara (Israel); 6) Shanidar (Iraq); 7) Teschik Tasch (Uzbekistan); 8) Denisova, Okladinov (Russian Siberia); Open Stars (other): 9) Hatnora (India) (Homo species indeterminate); 10)
Ngandong (H. erectus, Middle Pleistocene). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)
African and non-African populations, the revised rate yields an Out
of Africa chronology in the range of 130e90 ka, suggesting that the
Levantine MIS 5 artefactual evidence represents either permanent
or repeated temporary occupation by modern humans (Scally and
Durbin, 2012: 748). Younger estimates of Out of Africa on the
basis of mtDNA data may, it is argued, derive from processes of later
gene flow and drift/selection and/or complex demographic factors
including bottlenecks (Scally and Durbin, 2012: 751).
5. Evaluating Out of Africa
The earliest fossils of H. sapiens are identified in East Africa at c.
195e160 ka (White et al., 2003; McDougall et al., 2005), though
craniometric studies of African forms described as our species show
considerable phenotypic diversity (Gunz et al., 2009). Fossil evidence thus suggests that the emergence of H. sapiens occurred
towards the end of MIS 7, just prior to the onset of the colder and
drier conditions of MIS 6. The return to these conditions led to an
expansion of deserts, likely creating barriers to dispersal out of
Africa of this new species. If speciation estimates are correct, this
therefore means that other than a brief window at the end of MIS 7,
there may have been no real opportunity for modern humans to
leave Africa until MIS 5 (though see below; Dennell, 2009: 469). At
this time, territorial ranges in Africa appear to have expanded in
association with environmental amelioration, and Middle Stone
Age sites seem to be relatively abundant and well distributed in
East Africa (Basell, 2008) and the Saharan belt in MIS 5 (Smith et al.,
2007; Hill, 2009), providing the basis for a dispersal both within
and out of Africa (Osborne et al., 2008; Drake et al., 2011). As
indicated, it is indeed in MIS 5 that we find the first evidence for
anatomically modern humans outside of Africa. Fossil evidence at
Skhul and Qafzeh indicates their presence in the Levant from perhaps as early as 130e120 ka (Grun et al., 2005; Shea, 2008), i.e., the
early part of MIS 5 (5e).
The implications of the Levantine evidence are hotly debated.
On the one hand, certain researchers describe the Levantine
occupation by H. sapiens in MIS 5 as localized and temporary, and
argue that these humans were not entirely cognitively modern.
Though citing occupation dates that span approximately 40e
20,000 years, Mellars (2006b) describes the Levantine occupation
as a ‘short-lived event’. The Levantine Mousterian, often characterized by centripetal Levallois reduction methods and a variety of
tool types (e.g. points, scrapers, notches, burins), is viewed as
archaic technology, and inadequate to the task of adaptive competition with Neanderthals, who are perceived to have replaced
modern humans in this region (Shea, 2003, 2008; Mellars, 2006b).
Others, however, note the likelihood that Neanderthals and modern humans overlapped for a significant time period in MIS 5 in the
Levant, and argue for elements of modern human behavior in spite
of the apparently ‘limited’ technology (Grun et al., 2005). The
recent Neanderthal genome findings (Green et al., 2010) may
support the notion of an overlap. Furthermore, the suggestion that
use of Middle Palaeolithic technologies reflects a less competitive
H. sapiens, which had to await full cognitive modernity and the
related development of prismatic blades and more advanced projectile weaponry to outcompete Neanderthals, would seem to
reflect a traditional bias in the interpretation of the material record.
The variable appearance of advanced lithic technologies among
Asian populations of H. sapiens indicates the strong likelihood that
stone tool technologies rather reflect particular adaptive responses
N. Boivin et al. / Quaternary International 300 (2013) 32e47
to environmental circumstances and demographic parameters. The
shift to Early Upper Palaeolithic technology in the Levant appears to
have been associated with a reduction in prey size (Shea, 2008),
suggesting that either demographic pressure (Stiner et al., 1999,
2000) or environmental change, rather than cognitive development, drove the appearance of new technologies. A demographic
argument has recently been put forward for the appearance of
Upper Palaeolithic technology and other elements of material culture in western Eurasia, in this case based around the idea that
population needs to be of a certain density to retain cultural
innovations (Powell et al., 2009). Available evidence suggests that
parallels to Upper Palaeolithic technology do not emerge in South
Asia until c. 35 ka, which is much too late to reflect an initial
expansion of modern humans, and here as well, demographic
arguments have been forwarded for the appearance of more
advanced technologies (Petraglia et al., 2009).
The first out of Africa exit by H. sapiens is seen as part of a
limited range expansion of North African human and faunal populations (Tchernov, 1992; Lahr and Foley, 1994; Rabinovich and
Tchernov, 1995; Tchernov, 1996). Mellars (2006b) and other
scholars assume that this is a localized human expansion based on
the absence of similar fossils further afield. Yet, as discussed earlier, the fossil record is clearly biased; the absence of modern
human fossils in Arabia for MIS 5 is meaningless given that fossils
of Homo do not appear in the regional record until the Holocene.
Meanwhile, a recent multivariate analysis of cranial morphometric
data suggests affinity between early modern humans from the
Levant and terminal Pleistocene/early Holocene human populations from Australasia that is potentially reflective of a successful MIS 5 exit (Schillaci, 2008). From a palaeoenvironmental
perspective, it is difficult to identify factors that might have prevented dispersals further afield. Dispersal barriers both into and
within the Arabian Peninsula are at a minimum during MIS 5,
especially in MIS 5e and MIS 5a, when humidity and rainfall were
higher (Vaks et al., 2007; Fleitmann and Matter, 2009; Preusser,
2009) and when terrestrial environments are marked by lakes
and river systems (Parker, 2009; Petit-Maire et al., 2010;
Rosenberg et al., 2011). Genetic (mtDNA) data suggest that the
dispersal from Africa to Arabia of at least one other primate, the
Hamadryas baboon, in MIS 5 (Fernandez, 2009), though we would
approach this chronology with caution in light of calibration and
other issues. Glacial/stadial episodes, like MIS 4 and most of MIS 6,
would have been characterized by reduced monsoon rainfall and
greatly expanded deserts (the Sahara, Arabia and Thar deserts),
with the desert barriers much more extensive and uninviting than
those of the present day. By contrast the interstadials, especially
the more pronounced upswings of MIS 5e and MIS 5a would have
seen reduced deserts, which would furthermore have been significantly impinged upon by richer vegetation zones.
2010; Ozkan et al., 2010). So it is not necessarily surprising to find
that as further research is conducted, the picture of Out of Africa
has become more complex. Continued research supports an origin
of modern humans in Africa, and both expansion out of Africa and
replacement of local populations. However, the apparent simplicity
of this model is tempered by plausible evidence for a structured
African population, multiple exits, archaic admixture and the
existence of an increasing number of late archaic survivals in Asia.
These considerations, together with data suggesting that archaeological sites in southern Asia correspond with modern humans
extending back earlier than appreciated, suggest the need to
explore other models for Out of Africa. Below we present arguments for an earlier and more complex dispersal pattern for
modern humans, drawing in particular on relevant palaeoclimatic,
vegetation, faunal and genetic data and a set of maps that reconstruct vegetation patterns across Eurasia in MIS 4 and MIS 5 (see
Figs. 2e5, Table 1, Supplementary material C).
Table 1
Summary of vegetation zones mapped in Figs. 2e5.
Vegetation zone
Resource expectations
1. Mediterranean vegetation.
Abundant fruits and nuts and seeds,
including wild almonds, acorns,
Pistacia.
Many edible grain resources, some nuts
and fruits as above. Large herbivore
herds.
2. Mediterranean park-steppe,
the classic ‘Fertile Crescent’
vegetation famous for wild
cereal progenitors such as
wheat and barley.
3. The dry steppe of the Middle
East and southern Central Asia.
4. Desert
5. Sub-Desert/Sahel vegetation,
generally of the SudanoSindhian floristic province
6. Riverine corridors: marshes and
gallery forests.
7. Tropical savannah/woodlandgrass mosaic, including the
tropical evergreen zones of
India.
8. Dry tropical woodland, including
dry deciduous zones of India.
6. Modeling environments and movements
Despite the popularity in recent years of an Out of Africa model
featuring a single exit, a date of c. 60 ka for a rapid dispersal, and the
complete replacement of archaic species, it is clear that this model
is in need of revision given new datasets from genetics, archaeology, and palaeoanthropology. This parallels conceptual developments in domestication studies, where genetics and archaeology
initially supported a rapid, localized model for agricultural origins,
but gradually came to recognise more complex and protracted
processes involving multiple domestication events for many crops
(Brown et al., 2009). Ancient DNA studies, both of domesticated
plants and animals, and humans, have also begun to reveal the
importance of recent demographic processes, and their ability to
mask earlier genetic patterns (Haak et al., 2005; Larson et al., 2007,
37
9. Moist tropical woodland and
grassland mosaic, in particular
that of eastern India and Ganges
plain in which Dipterocarpaceae
including Shorea are frequent;
this zone extends to Southeast
Asia.
10. Moist tropical woodlands,
including both moist deciduous
zones like those in India and
true tropical rainforests.
Some edible seeds, but generally lower
value and higher effort than above, and
some tubers. Herbivore herds smaller
than above.
Minimal edible plant and animal
resources. No/low human
populations expected.
Small seeded grasses and forbs occur,
while seeds and fruit of many trees and
shrubs are available.
Herds of herbivores occur but these are
often of smaller herd size and/or body
size than for zones 2 or 7.
Wild grains of grasses, sedges, and
other aquatics as well as tubers are
frequent. Small game, birds and fish
are readily available, with some
larger game, especially in dry season.
Wild grass grains and other seeds are
numerous, including wild progenitors
of most tropical cereals; some tubers
are available and numerous edible
seasonal fruits. Large herbivore herds.
Numerous fruits, nuts and tubers.
Edible seeds of herbaceous plants
include wild pulses. Large herbivores
are frequent but in small groups.
Smaller game present but may be
hard to catch.
Wild grass grains and other seeds are
numerous, including seasonal wild rice
in local wetland; some tubers are
available and numerous edible seasonal
fruits. Large herbivore herds. Smaller
game present but may be hard to catch.
Wild seeds are sparse in time and space;
tree-nuts and fruits may be locally and
seasonally abundant but are likely to be
unpredictable in time and space. Larger
game is rare and smaller game is harder
to catch. Very low human population
densities expected.
(continued on next page)
38
N. Boivin et al. / Quaternary International 300 (2013) 32e47
Table 1 (continued )
Vegetation zone
Resource expectations
11. Tropical montane vegetation.
Some edible fruits, seeds and tubers,
but generally only localized and
seasonal. Few large game, and hard
to catch small game. Low human
population densities expected.
Many seasonal edible fruits and nuts,
including acorns. Tubers occur and
some edible grains and seeds. Large
herbivores are frequent but in small
groups; smaller game present but
may be hard to catch.
Few seasonal nuts and fruits. Minimal
seed or tuber resources. Sparse smaller
game. Low human population density
expected.
Many seasonal edible fruits and nuts,
including acorns. Some edible seeds,
but tubers rarer than in zone 12. Large
herbivores are frequent but in small
groups; smaller game present but
may be hard to catch.
Seasonal edible fruits and nuts, but
fewer than in zone 14, and with higher
toxicity. Edible seeds, seasonal and
localized. Large herbivores frequent,
as well as hard to catch small game.
12. Warm temperate hill and
sub-montane vegetation,
including the vegetation of
the Himalaya foothills, and
the evergreen broadleaf forests
of southern China.
13. High elevation coniferous
forests and desert/the Tibetan
plateau.
14. Mixed evergreen-deciduous
(mesophytic) forests.
15. Mixed conifer and temperate
deciduous
Meher-Homji, 2001). Despite this, many quantitative models of
past vegetation essentially flatten the earth and focus on temperature and rainfall in relation to latitude and longitude (e.g., Adams
and Faure, 1997; Prentice et al., 2000). For such quantitative models, the diversity of vegetation zones is greatly simplified even for
modern baseline maps. We have avoided this here through a less
quantitative, if admittedly more subjective approach, drawing
vegetation maps that take into account regional and topographic
variations evidenced in modern maps. In addition, we have taken
into account modern disjunct distributions of species or sister
species, since such disjunction imply past periods of more or less
continuous distributions between two regions, such as between
northeast of India and Sri Lanka or the savannas of Africa and India
(Asouti and Fuller, 2008: 72e73). Such disjunctions have long been
recognized in plant geography classifications, such as the “NuboSindian” and the “Eritreo-Arabian” floristic territories (e.g., Zohary,
1973) or recognition of the “Indo-African element” in the flora of
India (e.g., Legris, 1963; Awasthi, 1995). The maps provided here
must therefore be regarded as initial hypotheses that require systematic testing, through both more palaeoenvironmental proxy
records and quantitative modeling. To our knowledge, such
reconstructions have not been previously attempted for these
periods and geographical scales.
6.1. Taking steps eastwards
Vegetation reconstructions have been made at the macro-scale,
based on examining broad patterns of modern vegetation maps and
relating these to available paleobotanical and palaeoclimatic evidence sources. Synoptic vegetation descriptions, even in the modern day, are idealized simplifications, although of heuristic value
in recognizing that vegetation structure and composition are
strongly associated with climatic parameters of temperature and
rainfall (Huggett, 1995). Climatic parameters are regionally and
locally affected by topography and geographical coordinates, as
detailed maps of modern vegetation make clear (e.g., Wang, 1961;
Entry into the Arabian Peninsula from Africa was possible by
two main routes (see Fig. 2). The Sahelian vegetation of grasses and
shrubs would have extended far northwards into the Sahara and
across much of Arabia. Such environments can be expected to have
extended across the regions bordering the Red Sea, from Sudan’s
Jebel Elba through Egypt’s Jebel Abu Harba to the higher ground in
the Sinai, providing both a corridor into the Mediterranean environments of the Levant and a potential land route around the Red
Sea (see also Fig. 3 in Petraglia et al., 2010). A Bab el Mandab route at
Fig. 2. Vegetation reconstruction for western Eurasia in MIS 5. See Table 1 for details of vegetation zones. Labelled sites are: (A) Qafzeh; (B) Es-Skhul; (C) Jubbah Palaeolake; (D)
Shi’bat Dihya 1 (MIS 3); (E) Jebel Faya; (F) Aybut; (G) 16R Dune; (H) Jwalapuram; (I) Site 50.
N. Boivin et al. / Quaternary International 300 (2013) 32e47
39
Fig. 3. Vegetation reconstruction for western Eurasia in MIS 4. See Table 1 for details of vegetation zones. Site labels are as for Fig. 2.
the southern end of the Red Sea was possible, with parallel vegetation zones existing on either side of this strait. Palaeoshoreline
reconstructions over the past 150 ka indicate that there were a
number of periods when short sea crossings (ranging from <5 km
to <15 km) across the Red Sea were possible, thus suggesting there
is no reason to invoke seaworthy boats or seafaring technology and
skills (Lambeck et al., 2012). Moreover, potential short sea crossings
correspond with favourable environmental conditions on the Arabian Peninsula, such as at the transition between MIS 6 and 5e (e.g.,
Armitage et al., 2011). Once outside Africa, movement beyond the
Levant or southern Arabian Peninsula was likely to have been relatively straightforward in MIS 5. The disjunct distribution of
a number of tree and shrub species including many food species
that humans might have relied upon across Africa, the Yemeni
Fig. 4. Vegetation reconstruction for eastern Eurasia in MIS 5. See Table 1 for details of vegetation zones. Labelled sites (continued from Figs. 2 and 3) are: (G) 16R Dune; (H)
Jwalapuram; (I) Site 50; (J) Liujiang; (K) Huanglong; (L) Niah; (M) Huon; (N) Malakunanja; (O) Nauwalabila.
40
N. Boivin et al. / Quaternary International 300 (2013) 32e47
Fig. 5. Vegetation reconstruction for eastern Eurasia in MIS 4. See Table 1 for details of vegetation zones. Site labels are as for Fig. 4.
highlands and South Asia suggests the likelihood that their ranges
have contracted and that they would have occurred in much more
frequent patches in MIS 5. This would have included taxa with
edible fruits, such as drumstick trees (Moringa spp.), desert dates
(Balanites aegyptiaca), sebesten plums (Cordia spp.), karira (Capparis decidua), and tamarind (Tamarindus indica) (see Asouti and
Fuller, 2008). In addition, a similar range of edible wild milletgrasses, wild legumes and, on forest margins, yams (especially
Dioscorea bulbifera) and other tubers (e.g., Arisaema spp.) are shared
across such distributions. Attractive fauna for hunting, such as
gazelles and ostriches, would also have occurred through these
savannah-Sahelian corridors. Thus humans adapted to the northern
savannah woodlands and the Sahel in Africa would have been able
to expand through familiar environments out of Africa and across
Arabia to the similar environments that framed a reduced Thar
Desert. Riverine systems would furthermore have flowed east
across much of Arabia, providing additional dispersal corridors
(Edgell, 2006; Petraglia, 2007, 2011; Petraglia et al., 2012a). Lakes
and rivers in such humid stages as MIS 5 were present in areas now
characterized as desert in Arabia (Parker, 2009; Petit-Maire et al.,
2010; Rosenberg et al., 2011) and India (Achyuthan et al., 2007).
Archaeologically, such MIS 5 dispersal routes are not yet clearly
supported by artefactual evidence. Though Middle Palaeolithic sites
are abundant in Arabia, the majority have been identified from
surface contexts and remain undated (Petraglia and Alsharekh,
2003; Groucutt and Petraglia, 2012). New information from
stratified archaeological sites, however, is beginning to emerge.
Jebel Faya rockshelter, situated about 55 km from the Persian Gulf,
has been dated to c. 125 ka (MIS 5e) (Armitage et al., 2011). The
recovery of cordiform handaxes, foliates, Levallois and discoidal
cores and retouched tools is argued to be reminiscent of East
African industries, implying the early movement of H. sapiens into
Arabia. The most convincing case for a movement from Africa
comes with the recovery of abundant ‘Nubian’ sites situated along
riverine drainages in the Dhofar region of Oman (Rose et al., 2011).
Here, a Nubian assemblage was dated to 106 ka (MIS 5c), suggesting
connection with H. sapiens in northeast Africa. Middle Palaeolithic
industries situated in northern Arabia, on the shores of the Jubbah
Palaeolake, date to MIS 5 and MIS 7 (Petraglia et al., 2011, 2012a).
The Jubbah Palaeolake occurs in the Nefud Desert (Fig. 2), indicating that hominins penetrated this marginal environment in an
ameliorated period, where phytolith studies have demonstrated
the presence of a grassland with some trees.
The existence of two potential routes from Africa to Arabia, and
the presence of several main dispersal corridors through the Arabian Peninsula, suggest the possibility for multiple exits and more
complex dispersal scenarios. Favourable conditions in the Arabian
Peninsula, especially in the north, may have increased the range
overlap between Neanderthal and modern human populations,
presenting opportunities for interbreeding (Fig. 1). The potential
range of Neanderthals in MIS 5 likely extended well into presentday Iran, Afghanistan and Pakistan, and possibly south into the
Arabian Peninsula as well (Supplementary material B).
In contrast, MIS 4 saw widespread expansion of the desert in
northern Africa and the Arabian Peninsula (Fig. 3). This would have
impeded exit out of Africa other than via the Bab el Mandab, where
coastal environments of unknown but potentially quite variable
quality would have been encountered. Attractive grassland environments would have been located considerably inland from the
coasts, and probably at high elevations, such as in the Yemeni
highlands and the hills of southwest and eastern Oman peninsula.
Relict populations appear to be present at Jebel Faya and Shi’bat
Dihya across MIS 4 and extending into MIS 3, in both the Persian
Gulf region and in the Yemeni uplands (Armitage et al., 2011;
Delagnes et al., 2012). Pollen data furthermore suggest that in drier
periods like MIS 4, there was a reduction in the mangrove environments that often support rich coastal resources. Thus, although
the lowered sea levels of MIS 4 would have made crossing the Bab
el Mandab (and Arabian/Persian Gulf) easier, the coastal deserts on
either side are likely to have presented few attractions to huntere
gatherers. This is not to say that exit was not possible out of Africa
in MIS 4 (see Field and Lahr, 2005; Field et al., 2007), only that it
N. Boivin et al. / Quaternary International 300 (2013) 32e47
seems far less likely, and dependent on a number of coastal environmental parameters whose existence is debatable.
6.2. Passage to India
Subsequent to dispersal out of Africa and across Arabia, palaeoenvironmental data suggest a number of potential routes that
H. sapiens may have followed into the Indian subcontinent (Fig. 2).
These can be simplified into three broad routes, recognizing that
both hybrid and multiple routes are also possible and indeed likely.
Perhaps the most feasible route is the Sahel (marked as 5 on Fig. 2)
vegetation corridor, a route through environments that would have
featured many seasonal seed foods and recurrent medium-sized
game like gazelle and ostrich. The many disjunct tree species
between the dry tropical woodlands of India and Africa imply that
these vegetation zones were connected through intervening Arabia
in wetter periods in the past. These disjuncts include important food
sources that could have contributed to subsistence (e.g., desert
dates: Balanites, drumstick trees: Moringa, tamarind: Tamarindus).
The Sahel corridor route may have necessitated a sea crossing at the
Strait of Hormuz to present-day southern Iran, although a route
across or around the Persian Gulf is also feasible (Rose, 2010). A
second possibility is that there was an expansion of H. sapiens from
the Levant, via the Tigris/Euphrates riverine corridor that would
eventually become a major trade conduit in the region. Such riverine
corridors are always rich in small game, seeds foods and tubers,
although they might require rather different adaptations than the
Sahelian corridor. A third, perhaps less likely route is via the IndoIranian plateau. This would have necessitated following the Mediterranean park-steppe (classic Fertile Crescent vegetation) into the
Zagros Mountains (see Nasab et al., 2013). This vegetation possibly
extended across parts of the Iranian plateau or down to coast, then
onto India. This is perhaps the least likely route for various reasons
involving: 1) topographic complexity; 2) the need to adapt to significantly different environments and novel flora; and 3) the high
potential that Neanderthals dominated this region (see Fig. 1,
Supplementary material B). The range overlap with Neanderthals is
potentially important in light of the evidence for Neanderthal gene
flow into H. sapiens. Both of the latter two routes, and possibly also
the first, would likely have entailed ecological overlap with Neanderthals. So it is possible that regions east and perhaps even south
of the Levant would have provided opportunities for admixture
between modern humans and Neanderthals.
With a greatly reduced Thar Desert throughout MIS 5, entering
the Indian subcontinent would have been relatively straightforward compared to an entry in MIS 4 (Petraglia et al., 2012b). The
Sahel corridor that originated in North Africa and extended along
the southern Arabian Peninsula, subsequently skirting the Iranian
and Makran coasts, also likely framed a reduced Thar desert,
intergrading into the lusher savanna woodlands of the Deccan
plateau. This Sahel corridor thus provides a key potential and
previously unrecognized dispersal route between Africa and South
Asia. The Thar Desert itself need not necessarily have been a barrier
in MIS 5; palaeoenvironmental evidence from the desert in MIS 5
indicates a high percentage of C4 plants, increased summer rains
and a strong monsoon (Achyuthan et al., 2007). And, indeed, the
region can be expected to have had numerous rivers, and perhaps
playa lakes at this time. Based on Early Holocene parallels, we
expect that while many playas were quite saline, a few lakes would
have been freshwater recharged by subsurface drainage to the east
that was increased under stronger monsoons.
Is there fossil or archaeological evidence to support the presence
of modern humans in South Asia in MIS 5? As indicated, fossils are
lacking and skeletal remains of modern humans for India and Sri
Lanka are relatively young, and almost certainly tell us nothing about
41
the date of arrival of H. sapiens to the subcontinent. Archaeological
evidence is more suggestive, particularly in light of recent findings.
The recovery of numerous Middle Palaeolithic sites in many basins of
peninsular India, together with stratified occurrences post-dating
Late Acheulean assemblages (Misra, 2001), indicates that hominins
were present in the region for a considerable period of time.
Archaeological excavations at Jwalapuram, in the Jurreru Valley of
southern India demonstrate that Middle Palaeolithic hominins were
present by c. 78 ka (Petraglia et al., 2007, 2012c). Analysis of cores from
the Jwalapuram localities indicates affinities with sub-Saharan African MSA assemblages produced by H. sapiens (Clarkson et al., 2012).
The morphology and technology of the Jwalapuram cores may be
distinguished from technologies manufactured by Neanderthals in
Southwest Asia and Europe. Though the technology remains poorly
described, the potential presence of Middle Palaeolithic assemblages
dated 130e109 ka in the 16R Dune in the Thar Desert (Achyuthan
et al., 2007; Singhvi et al., 2010) and c. 125e75 ka in the Iranamadu
Formation of Sri Lanka (Deraniyagala, 1992; Abeyratne, 1996) also
suggest MIS 5 occupations that may represent H. sapiens, although of
course an archaic species cannot be ruled out (Petraglia et al., 2010).
While lithic technology does not provide a straightforward or definitive signature of modern human presence, in this case it does appear
to necessitate that we consider the possibility that H. sapiens was
present in South Asia by MIS 5a, if not earlier.
Palaeoenvironmentally, India appears to have been amenable to
human habitation in MIS 5 and into MIS 4 (Fig. 2). India had
favourable environments, such as broad areas of savannah (7) and
dry tropical woodlands (8), along with moist tropical woodland and
grassland mosaic (9) vegetation in the Ganges region. All of these
environments support a diversity of bovids and cervids, larger
herbivores (e.g., elephant, rhino), and numerous food plants. They
may also be regarded as quite well-buffered, in the sense that climatic oscillations would have opened woodlands and shifted but
not removed resources. While climatic fluctuation might have
extirpated local populations across much of more marginal environments like Arabia, populations in India may have been able to
persist through such oscillations. The inherent ‘habitability’ of
India, combined with a possible long-term modern human presence in the region, may go some way towards explaining the evidence for higher long-term population size in India than in
neighbouring regions (Atkinson et al., 2008; Petraglia et al., 2009).
If there was indeed an entry of H. sapiens into India in MIS 5, it is
possible, indeed probable, that modern humans encountered
archaic species in the region. A cranial fossil find from the Narmada
River valley supports the presence of an archaic human species
(initially identified as Homo erectus or Homo heidelbergensis, now
viewed as an indeterminate species of Homo) in the region sometime in MIS 6 or 5 (Athreya, 2007; Patnaik et al., 2009). Acheulean
sites are certainly present in the early to middle stages of the Middle
Pleistocene, indicating the long-term presence in the region of one
or more archaic species (Petraglia, 2010). Recent dating of Late
Acheulean sites to c. 140e130 ka in the Son valley of northern India
(Haslam et al., 2011) demonstrates that at least one archaic species
of Homo continued to inhabit India at the MIS 6/5 transition. The
coincidence with the earliest potential dates for Middle Palaeolithic
sites in the Thar Desert at 130e109 ka is obviously suggestive of an
early human presence. It is possible that new competition from
recently arrived anatomically modern humans led or contributed to
archaic extinctions at this time. Admixture with H. sapiens prior to
the extinction of archaic hominins cannot be ruled out.
6.3. Expansion into East and Southeast Asia
In many ways, the Indian subcontinent demonstrates significant
ecological similarity in comparison with the environments of
42
N. Boivin et al. / Quaternary International 300 (2013) 32e47
Africa. India is dominated by savannah and Sahel type vegetation,
as well as vegetation zones whose differences from these are not
radical, and similar food resources are attested to by disjunct taxa.
The region just east of India, however, features significantly different environments that may have posed new challenges to dispersing humans (Fig. 4). The northeast Indian region of Assam and
adjacent areas are characterized by high hills with dense tropical
forests, and seasonally flooded and swampy river valleys. Tropical
rainforests generally support low human population densities (cf.
Bailey and Headland, 1991) and necessitate specific rainforest
adaptations (e.g., techniques for hunting hard to catch arboreal
game such as monkey, see Piper et al., 2008; Perera et al., 2011).
Swamps may have supported lower populations due to higher
disease loads (cf. Sattenspiel, 2000), and if not true barriers would
nonetheless have reduced local populations. The impression of a
major barrier between the regions of India and Indochina is reinforced by biogeographical and genetic studies suggesting that a
wide range of animals are differentiated or speciated across this
region. A number of species have their range limits at the Brahmaputra River, including various species of mongoose (Herpestes
edwardsii, and the crab-eating mongoose, Herpestes urva), the
small-toothed palm civet, the large Indian civet, the hog-badger,
the Bengal fox, the Asian elephant, the pygmy hog, the hispid
hare, and various closely related primate species (e.g., the golden
langur, the hoolock gibbon and the stump-tailed macaque) (Corbet
and Hill, 1992; Vidya et al., 2005; Veron et al., 2007). The Brahmaputra also corresponds to four bird sister-species pair boundaries (Ripley and Beehler, 1990). The Brahmaputra itself, however, is
probably less as a physical barrier to gene flow than a marker
between two significantly distinct ecotones that are the true barriers to genetic exchange (Tosi, 2007). Other species differences
correlate to a region east of the Brahmaputra, in present-day
Myanmar, suggesting that biogeographic barriers in this region
are multiple. Examples of species separated by a Burmese biogeographic barrier include the mongoose species Herpestes javanicus
and Herpestes auropunctatus, and three bird sister-species pairs
(Ripley and Beehler, 1990). Other examples are the larger mammals
that are restricted to Southeast Asia, such as orangutans, gibbons,
sun bears and tapirs (and also the pandas of South China); none of
these ever entered South Asia, just as horses, camels and giraffids
never entered Southeast Asia from South Asia. This pattern of
barriers, along with a range of important human genetic (Metspalu
et al., 2004), cultural (e.g., milk drinking, Simoons, 1970), technological (e.g., the Movius Line, Movius, 1948) and domestic crop/
animal traits (Londo et al., 2006; Fuller, 2007; Larson et al., 2010)
that are or have also been differentiated across the Brahmaputra/
Burma boundary, suggests that it has long constituted one of the
most significant terrestrial biogeographic barriers of the Old World.
Accordingly, while it is not beyond the realm of possibility that
East and Southeast Asia will prove to have been colonized prior to
MIS 4, it would not be surprising to find that modern human colonization east of the Brahmaputra/Burma region is delayed, perhaps even until vegetation patterns change significantly in MIS 4.
While H. sapiens obviously displays significant behavioural flexibility and complexity, novel, more densely forested tropical landscapes of the type encountered in Burma would likely have
required significant new behavioural adaptations and toolkits.
Factors such as these may have discouraged expansion eastwards
unless population pressure arose or environments changed.
Archaeological evidence to date seems to support this hypothesis.
Technological differences between India and Indochina/Southeast
Asia are significant throughout the Pleistocene. The colonization of
Australia does not appear, on current evidence, to date to earlier
than 60 ka. Perhaps also significant is the fact that an increasing
number of archaic species of Homo appear to have survived until
relatively late in East and Southeast Asia. These include Homo
floresiensis, which survived on Flores until perhaps 17 ka
(Westaway et al., 2009), a possible Upper Pleistocene archaic
population in South China (Curnoe et al., 2012) and an as yet
unknown new hominin species from southern Siberia, which
existed as late as 50e30 ka (Krause et al., 2010).
If expansion east from India was significantly slowed by ecotone
change, then the opening up of this landscape in MIS 4 (Fig. 5) may
have had a significant impact on H. sapiens dispersal patterns. Bird
et al. (2004) note that in the tropics, glacial periods (and periods of
transition between interglacial and glacial periods) may have been
times of radiation, while interglacial periods were times of consolidation. Palaeoenvironmental data provide evidence that a corridor from east India through to insular Southeast Asia effectively
opened up at the beginning of MIS 4, as the moist tropical woodlands of western Indochina transitioned into a moist tropical
woodland and grassland mosaic landscape. Further east, this corridor would likely have opened onto a broad tropical savannah/
woodland-grass mosaic stretching across much of mainland
Southeast Asia. This has already been referred to as a ‘savannah
corridor’ and has been suggested to have constituted a key dispersal route into the region for modern humans (O’Connor and
Veth, 2000; O’Connor et al., 2001; Bird et al., 2004, 2005;
O’Connor, 2007; Wurster et al., 2010). The presence of H. sapiens in
Laos by MIS 3 is secure (Demeter et al., 2012), and although dating
is a problem, populations may be present in MIS 4. More open
environments may therefore have enabled rapid colonization of the
region down to the end of the Sunda plate. Effective dispersal
beyond this point may have had to await the evolution of an
appropriate maritime adaptation. On the other hand, though entry
into Sahul is often taken to have occurred no earlier than 45 ka
(O’Connell and Allen, 2004), occupations on the Huon peninsula
and in northern Australia (e.g., Malakunanja, Nauwalabila) may be
as old as 60e50 ka (Roberts et al., 1994), and in the Philippines as
early as 67 ka (Mijares et al., 2010).
Patterns of archaeological, palaeoenvironmental and other data
for China are complex and are beyond the scope of this article. We
would only note that the development of the Upper Palaeolithic is
young (35e30 ka) (Qu et al., 2012) and the earliest unambiguous
presence of H. sapiens in China appears to be no older than c. 40 ka
based on existing fossil and archaeological evidence (Shang et al.,
2010; Norton and Jin, 2009), though there are potential hints of an
earlier presence of H. sapiens fossils at sites such as Huanglong Cave
(Liu et al., 2010a), Liujiang (Shen et al., 2002; Wu, 2004) and Zhirendong (Liu et al., 2010b). Much of China is dominated by broadleaf
woodlands, rich in tree nuts and some tubers, many of which resources extend into the Himalayan foothills. While acorns and other
nuts are potentially rich resources, they would have necessitated the
development of cultural techniques of processing, which could have
contributed to a delay in the colonization of East Asia. Genetically, a
degree of Asia-based interbreeding between newly arriving modern
humans and archaic species is, as we have seen, suggested by data
from certain genes in the nuclear genome of contemporary humans
(Harding et al., 1997; Cox et al., 2008; Wall et al., 2009).
7. Discussion and conclusion
The question of whether the earliest H. sapiens fossils found in
the Levant reflect a failed dispersal of modern humans, or just the
first step in a journey that brought them as far as India by the end of
MIS 5, remains an open one. While mtDNA coalescence ages of 70e
50 ka are typically cited for the timing of modern human
movement out of Africa, the evidence in support of such a model,
along with such linked elements as the notion of a single exit and
rapid coastal dispersal, is actually minimal. Patterns in the fossil,
N. Boivin et al. / Quaternary International 300 (2013) 32e47
archaeological and genetic data do not conclusively support a c.
60 ka date for Out of Africa, and other readings of the evidence,
as well as other models, are possible. Here we have presented a
new analysis of the available evidence, together with a novel
attempt to model Pleistocene environments on the basis of
botanical
vegetation
reconstruction
techniques.
These
demonstrate that an MIS 5 exit is equally if not more plausible
than the MIS 3 or 4 exit that is currently widely advocated for
Out of Africa (see also Hetherington and Reid, 2010). In addition,
our analysis suggests that more complex scenarios for the dispersal
of modern humans should be favoured. These potentially involved
a structured African population of H. sapiens, multiple exits, multiple routes of dispersal, and a degree of interbreeding with archaic
species of Homo as anatomically modern humans moved eastwards. Given the current ambiguity of much of the evidence for
modern human dispersals out of Africa, it is critical that researchers
remain open to alternative readings of the available data. We have
focused here on the possibility that the modern human exit
recorded by fossil evidence in the Levant in MIS 5 does not represent a failed dispersal, and that in fact our species was not only in
the Levant but also the Arabian Peninsula during this marine isotope stage, and spread to India before the Toba eruption at 74 ka
(Petraglia et al., 2007). Another valid hypothesis we do not explore
here is that H. sapiens was able to leave Africa in MIS 6 via a
grassland corridor (Frumkin et al., 2011; see also Scally and Durbin,
2012). Yet another is that our species dispersed out of Africa shortly
after its first appearance c. 195 ka, in MIS 7 (Dennell and Roebroeks,
2005: 1102). One other possibility is that there were several, separate dispersals of our species out of Africa (Dennell and Petraglia,
2012). At the same time, we acknowledge that major demographic
changes occurred in MIS 4 and MIS 3, perhaps explaining the relatively young mtDNA coalescence age in living populations. The
increasing evidence for complexity as well as the clear patterns of
bias for all records, whether archaeological, fossil or genetic, suggests the need for an open mind to multiple scenarios for Out of
Africa, as well as for more rather than less complex models of
H. sapiens dispersal across Eurasia.
Acknowledgements
The field and analytical studies from which this article has
emerged have been funded by the British Academy, the European
Research Council (no. 295719), the Leakey Foundation, the Leverhulme Trust, the National Geographic Society, the Natural Environment Research Council, and the Society for South Asian Studies.
We thank three anonymous reviewers for helping us to make
improvements on this article and we are grateful to Michael Haslam for assisting with input of some references. We thank James
Blinkhorn and Huw Groucutt for inviting us to the Oxford conference and for their close reading of the text.
Appendix A. Supplementary materials
Supplementary material associated with this article can be
found in the online version, at http://dx.doi.org/10.1016/j.quaint.
2013.01.008.
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