R ES E A RC H
◥ on a somewhat regular basis. New finds of
REVIEW SUMMARY
PALEOANTHROPOLOGY
On the origin of modern humans:
Asian perspectives
Christopher J. Bae,* Katerina Douka,* Michael D. Petraglia*
BACKGROUND: The earliest fossils of Homo
sapiens are located in Africa and dated to the late
Middle Pleistocene. At some point later, modern
humans dispersed into Asia and reached the far-
away locales of Europe, Australia, and eventually
the Americas. Given that Neandertals, Denisovans,
mid-Pleistocene Homo, and H. floresiensis were
present in Asia before the appearance of mod-
ern humans, the timing and nature of the spread
of modern humans across Eurasia continue to
be subjects of intense debate. For instance, did
modern humans replace the indigenous popu-
lations when moving into new regions? Alterna-
tively, did population contact and interbreeding
occur regularly? In terms of behavior, did tech-
nological innovations and symbolism facilitate
dispersals of modern humans? For example, it
is often assumed that only modern humans were
capable of using watercraft and navigating to
distant locations such as Australia and the
Japanese archipelago—destinations that would
not have been visible to the naked eye from the
Map of sites with ages
and postulated early
and later pathways
associated with modern
humans dispersing
across Asia during the
Late Pleistocene.
Regions of assumed
genetic admixture are
also shown. ka, thousand
years ago.
Bae et al., Science 358, 1269 (2017) 8 December 2017
departure points, even during glacial stages
when sea levels would have been much lower.
Moreover, what role did major climatic fluc-
tuations and environmental events (e.g., the
Toba volcanic super-eruption) play in the dis-
persal of modern humans across Asia? Did ex-
tirpations of groups occur regularly, and did
extinctions of populations take place? Questions
such as these are paramount in understanding
hominin evolution and Late Pleistocene Asian
paleoanthropology.
ADVANCES: An increasing number of multi-
disciplinary field and laboratory projects focused
on archaeological sites and fossil localities from
different areas of Asia are producing impor-
tant findings, allowing researchers to address
key evolutionary questions that have long per-
plexed the field. For instance, technological
advances have increased our ability to success-
fully collect ancient DNA from hominin fossils,
providing proof that interbreeding occurred
H. sapiens fossils, with increasingly secure dating
associations, are emerging in different areas
of Asia, some seemingly from the first half of
the Late Pleistocene. Cultural variability dis-
cerned from archaeological studies indicates
that modern human behaviors did not simply
◥
spread across Asia in a
ON OUR WEBSITE
Read the full article
time-transgressive pat-
tern. This regional varia-
at http://dx.doi. tion, which is particularly
org/10.1126/ distinct in Southeast Asia,
science.aai9067 could be related at least
..................................................
in part to environmental
and ecological variation (e.g., Palearctic versus
Oriental biogeographic zones).
OUTLOOK: Recent findings from archaeology,
hominin paleontology, geochronology, and ge-
netics indicate that the strict “out of Africa”
model, which posits that there was only a single
Downloaded from http://science.sciencemag.org/ on April 2, 2020
dispersal into Eurasia at ~60,000 years ago, is
in need of revision. In particular, a multiple-
dispersal model, perhaps beginning at the
advent of the Late Pleistocene, needs to be ex-
amined more closely. An increasingly robust
record from Late Pleistocene Asian paleoan-
thropology is helping to build and establish
new views about the origin and dispersal of
modern humans.
▪
The list of author affiliations can be found in the full article online.
*Corresponding author. Email: cjbae@hawaii.edu (C.J.B.);
douka@shh.mpg.de (K.D.); petraglia@shh.mpg.de (M.D.P.)
Cite this article as C. J. Bae et al., Science 358, eaai9067
(2017). DOI: 10.1126/science.aai9067
1 of 1
R ES E A RC H

◥ cifically, how the Asian record contributes to ad-

REVIEW dressing such questions.

The big questions

PALEOANTHROPOLOGY Findings from archaeology, hominin paleontology,
geochronology, genetics, and paleoclimatology

On the origin of modern humans: have all been contributing to a better understand-
ing of the Late Pleistocene human evolutionary
record in Asia. Here we discuss some of the big
Asian perspectives questions that paleoanthropologists are investi-
gating across Asia: Can modern human dispersal
out of Africa be considered a single event occurring
Christopher J. Bae,1* Katerina Douka,2,3* Michael D. Petraglia2,4*
only after 60 ka, or is the picture more compli-
cated? By which route(s) did modern humans dis-
The traditional “out of Africa” model, which posits a dispersal of modern Homo sapiens
perse across Asia? What was the nature of the
across Eurasia as a single wave at ~60,000 years ago and the subsequent replacement of
interactions between modern humans and hominin
all indigenous populations, is in need of revision. Recent discoveries from archaeology,
groups already present in Asia? What role did
hominin paleontology, geochronology, genetics, and paleoenvironmental studies have
geographic and/or paleoenvironmental variations
contributed to a better understanding of the Late Pleistocene record in Asia. Important
play in modern human dispersals?
findings highlighted here include growing evidence for multiple dispersals predating
60,000 years ago in regions such as southern and eastern Asia. Modern humans moving into Can the modern human dispersal be
Asia met Neandertals, Denisovans, mid-Pleistocene Homo, and possibly H. floresiensis, with considered a single event occurring only

Downloaded from http://science.sciencemag.org/ on April 2, 2020

some degree of interbreeding occurring.These early human dispersals, which left at least some after 60 ka?
genetic traces in modern populations, indicate that later replacements were not wholesale.
Variations of the “out of Africa” (OoA) model may

T
he origin of modern humans has long per-
plexed us. As the well-known paleoanthro-
pologist William Howells remarked more
than four decades ago [(1), p. 477], “That
part of human history covering the emer-
gence of modern man and his regional differ-
entiation continues to be surprisingly obscure.
Locations of some elements of agreement or con-
troversy…have long been clear, but the dimensions
of the whole problem are far from obvious. The
trees are familiar, but the forest is not.”
Perhaps the primary reason for this is that
there has been no universal consensus on the traits
that make us human. Arguments for a large cra-
nial capacity defining our species went by the
wayside with the realization that Neandertals,
whose cranial capacity is slightly larger than ours
on average, were not actually Cossack soldiers
or pathological anomalies, but rather were of
a much greater geological age than previously
thought and in fact were penecontemporaneous
with the earliest modern humans (2, 3). Further,
we once assumed that “culture” clearly distin-
guished us from all other life forms. However,
when Jane Goodall (4) returned from her field
studies in the 1960s with the discovery that chim-
panzees made and used tools, it raised questions
about whether we could continue to use a broad
variable such as culture to define humans. The
few generally agreed-upon autapomorphies spe-
cific to modern humans include such phenotypic
traits as the presence of a mental eminence and a
globular braincase. However, given the absence
1
Department of Anthropology, University of Hawai‘i at Manoa,
2424 Maile Way, 346 Saunders Hall, Honolulu, HI 96822, USA.
2
Max Planck Institute for the Science of Human History, Kahlaische
Strasse 10, D-07743 Jena, Germany. 3Research Laboratory for
Archaeology and the History of Art, School of Archaeology,
University of Oxford, 1-2 South Parks Road, OX1 3TG Oxford,
UK. 4Human Origins Program, National Museum of Natural
History, Smithsonian Institution, Washington, DC 20560, USA.
*Corresponding author. Email: cjbae@hawaii.edu (C.J.B.);
douka@shh.mpg.de (K.D.); petraglia@shh.mpg.de (M.D.P.)
of the latter trait among the recently reported
early modern human fossils from Jebel Irhoud,
Morocco (5, 6 ), it may be possible that it devel-
oped much more recently, only within the past
130,000 years.
Some of the earliest morphologically modern
humans are reported from the sites Omo Kibish,
dating to ~195 thousand years ago (ka) (7), and
Herto, dating to ~160 ka (8), both located in the
Horn of Africa (Fig. 1A). These fossils have been
used for the past several decades to support an
East African origin for modern humans, corrobo-
rating similar findings from genetic studies (9–11).
Yet the recent discovery of modern human fossils
from Jebel Irhoud dating to ~310 ka (5, 12) raises
important questions about the singular role of
East Africa in the genesis of modern human mor-
phology. Further, questions exist about whether
so-called modern human behaviors appeared
around the same time as modern human mor-
phology, with most studies supporting a slower
behavioral transition that occurred over the span
of several hundred thousand years in Africa (13).
Despite the general acceptance that Homo
sapiens arose in Africa, the initial arrival and
survival of modern humans in different areas of
the world continue to be strongly debated. Over
the past several decades, Asia has been receiv-
ing increasing attention in these discussions,
particularly because it is considered the conduit
through which H. sapiens arrived in distant lo-
cales such as Western Europe, Australia, and
eventually the Americas. Importantly, the Asian
continent, bounded roughly by the Pacific, Indian,
and Arctic Oceans on three sides and Europe to
the west, includes a wide range of latitudinal,
longitudinal, and even altitudinal variation, which
has major implications for human evolution (14).
Because the questions of what, where, how, and
especially why with regard to our becoming “hu-
man” continue to be of great interest, we evaluate
the debate on modern human origins and, spe-
be broadly categorized as follows: (i) a single dis-
persal occurring during marine isotope stage
(MIS) 5; (ii) multiple dispersals beginning dur-
ing MIS 5; (iii) a single dispersal occurring dur-
ing MIS 3; and (iv) multiple dispersals beginning
during MIS 3. We detail each of these broadly de-
fined models below, but we begin with the single-
dispersal MIS 3 iteration because it has received
the greatest attention, followed by the multiple-
dispersal MIS 5 model.
Single dispersal during MIS 3
The traditional OoA model proposes that a single
major dispersal event of modern humans out of
Africa and into Eurasia occurred some time after
60 to 50 ka (15). In this model, earlier dispersals
by anatomically modern humans into the Levant
(e.g., Qafzeh and Skhul) were minor in scale and,
for all intents and purposes, evolutionary dead
ends. This OoA model was largely supported by
early genetic studies and to various extents by
archaeology, geochronology, and hominin pale-
ontology (16–21). The 60-ka dispersal continues to
be based primarily on genetic studies of present-
day human population variation across the world.
Given the degree of variability in the genetic clock
(because of varying mutation rates and other un-
certainties), it may be possible that this event
occurred earlier or later than the 60-ka marker
(18, 22). More recent whole-genome studies (23–25)
suggest that a single major dispersal event oc-
curred during which “all contemporary non-
Africans branched off from a single ancestral
population” [(26), p. 179] some time during the
Late Pleistocene (27 ). Archaeologists have also
set the boundary between 60 and 50 ka on the
basis of the timing of the behavioral and tech-
nological transition from the Middle to the Upper
Paleolithic (20, 21, 28).
Multiple dispersals during MIS 5
Another dispersal scenario that is receiving
increasing attention is the possibility that multiple

Bae et al., Science 358, eaai9067 (2017) 8 December 2017 1 of 7

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dispersals began by the beginning of the Late
Pleistocene (14, 29–31). Given the increasing num-
ber of paleontological and archaeological reports
of hominins from sites in different areas of Asia
that supposedly predate 60 ka, it would appear
that earlier dispersals out of Africa made it to the
Levant (Q afzeh and Skhul) and probably also
to South, East, and Southeast Asia (Fig. 1A and
Table 1) (32–42). At least some traces of these
earlier dispersals would be expected to be pres-
ent in the modern record. A recent genetic study
seems to support the fossil and archaeological
records in that ~2% of the genome of modern
Papua New Guineans derives from an ancestry
that predates the postulated 60-ka expansion by
modern humans out of Africa (43).
Single dispersal during MIS 5 and multiple
dispersals during MIS 3
These two models receive the least support from
the various scientific records. The model of a sin-
gle dispersal during MIS 5 only works if modern
ers have hypothesized two distinct paths: The
first involves crossing over from northern Egypt
to the Sinai Peninsula, whereas the second in-
volves crossing the Bab al Mandab Strait to
Yemen in the southernmost part of the Arabian
Peninsula (30, 44–46) (Fig. 1A). The Bab al Mandab
Strait is usually about 20 km wide, and during
glacial periods, it may have only been about 5
to 15 km wide. Given the short distances in-
volved, the other continent would have been vis-
ible from the points of both departure and arrival
(47), as is the case today. Yet this route requires a
water crossing that would have made rafting a
necessity. The Sinai Peninsula, on the other hand,
is the only land corridor that has persisted since
the permanent closing of the Tethys Seaway during
the Miocene. This land route remains the stron-
gest, and, for some, the most parsimonious can-
didate for the key dispersal pathway beyond
Africa (48, 49).
There is little consensus as to the route(s)
taken once outside of Africa. Two major direc-
The basic tenet of the southern dispersal hy-
pothesis is an eastward expansion from the Horn
of Africa via the Strait of Bab al Mandab, across
the southern part of the Arabian Peninsula, along
the Yemen-Oman littorals to the Persian Gulf, and
then along the coast of the Indian subcontinent
(18). Upon arrival at the Sunda Shelf, and after a
short hiatus, humans dispersed to Sahul, eventu-
ally reaching Australasia soon after 60 ka (53). The
southern coastal dispersal route out of Africa and
around the Indian Ocean continues to receive sub-
stantial attention, in part because a number of
genetic studies seem to support it [e.g., (18, 19, 25)].
However, for modern humans to have been able to
traverse the southern coastal route, regular access
to fresh water and utilization of marine resources
would have been necessary (54, 55). Unfortunately,
clear evidence for a coastal dispersal around the
Indian Ocean is not present for regions outside
Sunda, and instead, dispersal from the Sinai
across terrestrial regions of Arabia and southern
Asia is surmised as the main route of population

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humans suddenly appeared all across Asia in large
numbers and remained so continuously through-
out the Late Pleistocene. Although a growing num-
ber of sites have been dated to MIS 5 and 4, and
a number of these are considered to be associated
with modern humans, the data do not seem to
reflect a major dispersal event at this early date
(compare Fig. 1, A and B). No genetic studies of
which we are aware support such a model. The
model of multiple dispersals during MIS 3 only
works if there is no evidence for modern humans
before MIS 3 (compare with Fig. 1A).
By which route(s) did modern humans
disperse across Asia?
The most widely discussed OoA routes have East
Africa as the departure point into Asia. Research-
tional dispersal models have been proposed:
northern and southern. The northern dispersal
model posits that humans moved north of the
Sinai and through the Negev region to reach the
eastern Mediterranean Levant (48). Evidence to
support this includes the early modern human
sites of Skhul and Qafzeh in Israel that date to be-
tween 120 and 90 ka and proposed technological
similarities between artifact assemblages in north-
eastern Africa and the Levant (44, 48, 50, 51).
After this period, there is no evidence for the
presence of modern humans in the Levant until
about 55 ka or later, based on the recent findings
from Manot Cave (52). Because of this paucity of
evidence, the early presence of modern humans
in the Levant is considered by many to be a failed
dispersal event.
movements (29–31). It is possible that although
coastlines provided favorable habitats, at least
occasionally, reliance on such resources should
be seen as only one component of the whole hu-
man subsistence package (31).
FADs and TAQs
An important point in developing a more con-
crete understanding of the timing and direction
of human movements outside Africa is that the
first appearance datum (FAD) of modern humans
can serve as terminus ante quem (TAQ) for when
these dispersals began. In this regard, with the
emphasis on the supposed “end points,” the key
arrival areas are Europe, Australasia, the Japanese
archipelago, and eventually the New World.
Europe has long been at the forefront of these

Fig. 1. Map of sites and postulated migratory pathways associated migration pathways of H. sapiens, based on evidence from the most important
with modern humans dispersing across Asia during the Late sites dated to between 60 and 30 ka. Yellow stripes and translucent
Pleistocene. Archaic human distributions are also shown. (A) Initial orange represent tentative ranges for Neandertals and early Homo sapiens,
dispersal(s) of Homo sapiens, based on sites predating 60 ka. (B) Subsequent respectively.

Bae et al., Science 358, eaai9067 (2017) 8 December 2017 2 of 7

R ES E A RC H | R E V IE W

discussions because it is a heavily studied region A major colonization event that has not re- admixture and the sharing of culture to competi-
and an area that was occupied by Neandertals ceived as much attention is the peopling of the tion between and possibly the extinction of one of
for several hundred thousand years, with only Japanese archipelago; it has been suggested that the populations (45). Data from genetics and ar-
about a 5000- to 3000-year overlap with modern this could have occurred during MIS 6, when a chaeology have directly contributed to increasing
humans after initial colonization by the latter land connection was likely present, or some time our understanding of what may have happened
group (56). Although migration corridors between during MIS 3, when no land connection existed when Late Pleistocene humans moved into new
Europe and the Levant may have been present (28, 65–67). Given that only a few sites in Japan regions of Asia.
during MIS 5 (57), evidence for an unequivocal predate 40 ka, and that there are questions about
FAD by modern humans before 44 ka is absent. the context and/or artifactual nature of the ma- Introgression
The earliest example of fully modern human terials at those localities, the earliest presence Over the past decade, technological advances in
anatomy in Europe is the mandibular fragment of modern humans in Japan is considered to be the field of ancient DNA analysis have allowed
from Kent’s Cavern in England and the Cavallo about 40 ka (67). Assuming that this date is cor- scientists to obtain uncontaminated genome-
teeth from southern Italy (58, 59), dating to be- rect, the initial colonization would have had to wide data from Pleistocene hominin fossils. This
tween 41 and 44 ka (Fig. 1B). However, given the involve some type of watercraft and a high degree has shown that a fair degree of introgression oc-
geographic location of both sites, and assuming of seafaring skill (28, 34, 65, 66). curred between Neandertals and modern humans
that dispersals into Europe originated from the The timing and route by which humans arrived (27, 72–74). In fact, estimates of Neanderthal DNA
Levant or elsewhere in northern Asia and traveled in the Americas have been the subjects of a long present in non-African modern humans generally
westward, this opens the possibility that earlier and often heated debate (68, 69). Most data ap- range between 1 and 4% (72). One estimate from
evidence for modern humans in central or eastern pear to support colonization through Beringia ei- the Romanian Pestera çu Oase 1 fossil is as high as
Europe may be present. ther via the ice corridor or the coastal route some 9%, suggesting that this particular modern human
It has long been argued that modern humans time during or right after the last glacial maxi- with a minimum age of 40,000 years may have
were the only hominin taxon capable of peopling mum (~20 to 15 ka) (68). However, to reach North had a Neandertal ancestor as recently as four to six

Downloaded from http://science.sciencemag.org/ on April 2, 2020

Australasia, particularly because it would have America, human foragers had to have first arrived generations back (75). The recent genetic iden-
involved the ability to build sturdy watercraft in Siberia. The earliest peopling of Siberia north tification of penecontemporaneous Denisovans
and navigate the open seas (60). The peopling of of 50°N appears to have only occurred some time in southern Siberia further complicates the Late
Australasia took on greater importance once it between 50 and 45 ka, as indicated by the human Pleistocene human evolutionary picture in Asia
was realized that it likely occurred some time femur from Ust’-Ishim in western Siberia (70), (76–79). Reich and colleagues (80) estimated that
between 60 and 40 ka (61, 62). The recent re- and the earliest peopling north of 60°N occurred about 5% of modern-day Melanesian DNA origi-
analysis of the Madjedbebe (Malakunanja II) site perhaps only as recently as 32 ka (68, 71). Nean- nates from this ancestral Denisovan population,
in northern Australia pushes the initial peopling dertals and Denisovans were clearly in southern although more recent estimates are between 1 and
of the region back to at least 59 ka and possibly Siberia during the first half of the Late Pleisto- 3% (27, 81). Further, Prüfer and colleagues (73)
65 ka (63), although no fossils have been re- cene, but more northward dispersals were likely found that Neandertals, who were also present at
covered from the site. With early modern human by modern humans (Fig. 1B). Denisova Cave, interbred with Denisovans; simi-
fossils being reported in mainland Southeast Asia larly, gene flow occurred between Denisovans
(36, 39, 40), Indonesia (42), and the Philippines How did modern humans interact with and a yet-to-be identified hominin population. In
(38) (Fig. 1A) that predate the findings from hominin groups already present in Asia? the latter case, Prüfer and colleagues (73) pos-
Madjedbebe (65 ka) and New Guinea [49 ka (64)], When humans arrive in a new territory, one of tulated that the gene flow could be from an an-
this is perhaps not all that surprising. several things may happen, generally ranging from cestral population, such as H. erectus. Thus, a

Table 1. Background data for all Asian Late Pleistocene sites associated with early modern humans that appear in Fig. 1A. Triple plus signs indicate
high confidence (hominin fossils and archaeology found in clear association with solid dates); double plus signs, medium confidence (hominin fossils and/or
archaeology found in association or questionable association with dates); single plus sign, weak confidence (hominin fossils and/or archaeology found in
questionable association with dates).

Site number in Fig. 1A Site Present-day country Proposed age range Hominin fossils Archaeology Confidence Reference
(thousand years)

9 Skhul, Qafzeh Israel 120 to 90 Yes (H. sapiens) Yes +++ (50, 51)
.....................................................................................................................................................................................................................................................................................................................................
10 Jebel Qattar Saudi Arabia 75 None reported Yes ++ (110)
.....................................................................................................................................................................................................................................................................................................................................
11 Mundafan Saudi Arabia 100 to 80 None reported Yes ++ (31)
.....................................................................................................................................................................................................................................................................................................................................
12 Aybut Al Auwal Oman 105 None reported Yes ++ (111)
.....................................................................................................................................................................................................................................................................................................................................
13 Jebel Faya C United Arab Emirates 125 None reported Yes ++ (112)
.....................................................................................................................................................................................................................................................................................................................................
14 Katoati, 16R Dune India 96, 80 None reported Yes ++ (95)
.....................................................................................................................................................................................................................................................................................................................................
15 Jwalapuram India 85 to 75 None reported Yes ++ (33)
.....................................................................................................................................................................................................................................................................................................................................
16 Huanglong China 100 to 80 Yes (H. sapiens) Yes ++ (35)
.....................................................................................................................................................................................................................................................................................................................................
17 Luna China 120 to 70 Yes (H. sapiens) Yes ++ (40)
.....................................................................................................................................................................................................................................................................................................................................
18 Liujiang China 130 to 70 Yes (H. sapiens) None reported + (32)
.....................................................................................................................................................................................................................................................................................................................................
19 Fuyan China 120 to 80 Yes (H. sapiens) None reported ++ (37, 41)
.....................................................................................................................................................................................................................................................................................................................................
20 Zhiren China 100 Yes (H. sapiens) None reported ++ (36)
.....................................................................................................................................................................................................................................................................................................................................
21 Tam Pa Ling Laos 63 to 46 Yes (H. sapiens) None reported ++ (39)
.....................................................................................................................................................................................................................................................................................................................................
22 Callao Philippines 67 Yes (H. sapiens?) Yes ++ (38)
.....................................................................................................................................................................................................................................................................................................................................
23 Lida Ajer Sumatra 73 to 63 Yes (H. sapiens) Yes ++ (42)
.....................................................................................................................................................................................................................................................................................................................................
24 Madjedbebe Australia 65 None reported Yes ++ (63)
.....................................................................................................................................................................................................................................................................................................................................

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R ES E A RC H | R E V IE W
growing number of studies indicate multiple admix-
ture events between modern humans, Neandertals,
Denisovans, and a nonidentified population—
events assumed to have occurred in Asia (11, 73, 82).
Substantial overlap in time ranges in these same
areas also lends support to likely interactions be-
tween these different populations (Fig. 2).
Genetic studies have shown that the admix-
ture between non-African modern humans and
Neandertals could have occurred as recently as
40 to 86 ka (70, 75). The younger date would
tend to support the 60-ka OoA model, whereas
the older admixture date could be interpreted
as one of the early dispersals thought to have
occurred during MIS 5. Malaspinas and colleagues
(23) presented an even more complex admix-
ture model and suggested a series of events that
occurred between 72 and 42 ka, which included
a “ghost” lineage. To further complicate the matter,
Posth and colleagues (74) recently suggested that
the beginning of H. sapiens–Neandertal intro-
gression could date to as far back as ~270 ka.
Although indications of introgression now
commonly appear in the genetic literature, we
should attempt to determine whether it is pos-
sible to identify how genetic interchange appears
phenotypically. A range of studies of hybrids among
closely related nonhuman primates identified ex-
amples of dysgenesis (“hybrid weakness”) and/or
heterosis (“hybrid vigor”) when evaluating a di-
versity of size and shape variables (83). A number
of fossils have been proposed as possible hybrids
in the Late Pleistocene human fossil record (84),
including from Pestera çu Oase in Romania (85)
and Zhirendong in China (36). Martinón-Torres
Central Asia
Annual precipiation West Asia
[mm]
MIS
290
330
250
1 10 ka
2 Kulbulak
25
Yafteh
Üçağızlı
3 Teshik Tash
Ksar Akil
50 Kebara
Manot
Shanidar
4 Callao
Amud
75
5 Qafzeh
100
Skhūl
Tabun
125
-40
-44
-36
-32
300
500
700
NGRIP East Asian monsoon Neandertals
δ18O [‰] Annual precipiation
[mm]
Fig. 2. Age ranges for the presence of different hominin taxa (mod-
ern humans, Neandertals, Denisovans, and H. floresiensis) in all
major regions of Asia. The dotted line represents the proposed 60-ka
Bae et al., Science 358, eaai9067 (2017) 8 December 2017
and colleagues (86) even recently suggested that
H. floresiensis may be a hybrid. The key to
phenotypically identifying a hybrid may be to
observe, with some regularity, unusual traits (e.g.,
supernumerary teeth) that are not present in
the proposed parent population but suddenly
appear in the supposed offspring.
Cultural diffusion
In a straightforward scenario, modern humans
would have moved out of Africa and into Asia
carrying with them a set of standard “modern”
behaviors (e.g., the use of blades, microblades,
art, and symbolism). This is commonly referred
to as the “human revolution” model, thought
to reflect the Middle to Late Paleolithic transi-
tion (21, 28, 55). Numerous studies, however,
have questioned whether a behavioral revolu-
tion actually took place and whether a model,
originally developed for the western European
record, is suitable to other regions of the Old
World [e.g., (13, 34, 87–90)]. Further, the human
behavioral and skeletal records do not line up
neatly: Modern human behaviors often appear
in areas where modern human fossils do not. Be-
haviors traditionally considered to be represent-
ative of modern humans now have been reported
in association with Neandertals, Denisovans, and
possibly other hominin taxa (91–93).
Blade technology, long considered one of the
core components of the Upper Paleolithic in
Asia, appeared in western Asia after 50 ka and
arrived in South and North Asia sometime after-
ward. Microblades appeared during early MIS 3
in South Asia and late MIS 3 in North Asia, be-
North Asia Central Asia
Mal’ta
Pokrovka
Ust’-Ishim
Okladnikov
Obi Rakhmat
Chagyrskaya
Denisova
Denisova
Modern humans
exodus out of Africa supported by various studies. If the 60-ka model is
correct, modern humans should not appear in Asia earlier than that.
NGRIP, North Greenland Ice Core Project.
coming more prominent after 35 and 25 ka, re-
spectively (31, 33, 34, 94, 95). Interestingly, early
blade and microblade technologies have yet to be
identified in Southeast Asia, including southern
China. Because Southeast Asia represents a dif-
ferent biogeographic zone (Oriental) than the
other regions (Palearctic), this suggests that dif-
ferent ecological demands required the develop-
ment of a different behavioral toolkit to survive
(14, 34, 96–98). Early evidence of rock art (99)
and deep-sea pelagic fishing (100) in Southeast
Asia are clear signs of other forms of modern
human adaptation.
It has long been thought that modern human
foraging groups moving over the northern route
through Asia carried with them a modern be-
havioral package (28, 34, 101), sometimes equated
with the Initial Upper Paleolithic technocomplex.
Interestingly, however, the recent identifica-
tions of Denisovans and Neandertals in Denisova
Cave—a site that has traditionally been known
for the presence of a diverse Upper Paleolithic
Downloaded from http://science.sciencemag.org/ on April 2, 2020
industry (e.g., blades, bone tools, and bone or-
naments) (102)—has complicated this dispersal
model, particularly given the apparent absence
of H. sapiens fossils at the site (Box 1). A set of
perforated teeth and ostrich eggshell and bone
pendants were excavated from layer 11 of Denisova
Cave, the same layer that is assigned to Nean-
dertals (sublayer 11.4) and Denisovans (sublayer
11.2) (73). This may add to the growing evidence
that, at least on a small scale, Neandertals and
perhaps other hominins were capable of sym-
bolic behavior (91, 92). Alternatively, we may
be witnessing a series of local extinction and/or
East Asia South Asia Southeast Asia
10 ka
Zhoukoudian 25
Upper Cave
Yamashita-Cho
Batadomba Niah Cave
Tianyuan
Fa Hien Tabon 50
Tam Pa Ling
75
Liang Bua
Huanglong
Zhiren
Luna 100
Fuyan
125
Denisovans H. floresiensis
4 of 7
R ES E A RC H | R E V IE W

However, increasing findings indicate that multiple

Box 1. Who were the Denisovans? dispersals out of Africa by early modern humans
On the basis of DNA sequencing of a juvenile finger bone found almost a decade ago, a began during MIS 5, resulting in their earlier ar-
previously unknown Late Pleistocene hominin population was identified from Denisova Cave in rival in distant localities in the Levant, South Asia,
the Altai Mountains in southern Siberia, now referred to as the Denisovans (76, 77). The Southeast Asia, and China (33, 35–37, 39–42, 50)
hominin fossil assemblage from Denisova Cave includes this distal manual phalanx from layer (Fig. 1A).
11.2 (Denisova 3) and an upper left M3 or M2 tooth from layer 11.1 (Denisova 4), found in The initial dispersals out of Africa during MIS
association with various archaeological materials typical of both the late Middle Paleolithic (e.g., 5 were likely by small groups of foragers, and
Levallois flakes) and the Upper Paleolithic (e.g., microblades and ornaments made from ground these appear to have moved along both a south-
stone) (113). In addition, an upper molar (Denisova 8) was found at the interface of layers 11.4 ern and a northern route. The initial dispersal
and 12 in 2010, tentatively also identified as an M3 or M2 (78), and a lower left molar (Denisova 2) by modern humans northward reached at least
was recently added to the small number of fossils from the site (79). The archaeological sequence Qafzeh and Skhul. The southern route may have
begins at about 250 ka; Denisova 8 has been tentatively dated to >50 ka, whereas Denisova 3 and followed the coast around the Indian Ocean
4 have been dated to between 50 and 30 ka (76–78), although it is likely that these dates will be (18–20), but archaeological evidence dominates
revised after further geochronological study. for inland dispersal corridors, where a diversity
Despite the fact that genetic analyses indicated that the Denisovans were statistically of habitats occur and where reliable freshwater
significantly different from both modern humans and Neandertals, comparative morphological rivers, lakes, and animal resources were present
studies of the phalanx and molars were inconclusive (11, 77, 78). Thus, these remains were (29–31). Sites dating to early MIS 5 to MIS 4 that
referred to simply as Denisovans and not assigned a new species name. Since these initial studies, appear in places such as India, southern China,
however, suggestions have arisen that Denisovans may be present in the fossils of Chinese mid- Laos, Sumatra, and the Philippines are all located
Pleistocene Homo (or even Penghu 1 from Taiwan) (14, 86, 114). Initial comparative studies have a fair distance from any paleocoasts. At least some
shown that the Denisovan dentition displays similarities with those of the Chinese Xujiayao hominin of these early dispersals left low-level genetic traces

Downloaded from http://science.sciencemag.org/ on April 2, 2020

fossils and Teshik-Tash and Oase 2 fossils from western Asia (115, 116). To further complicate
matters, at least three Neandertal fossils were also identified at the site (Denisova 5, 9, and 11)
(73, 117).
replacement events at Denisova Cave that in-
volved all three hominin populations, with modern
humans solely responsible for the Upper Paleo-
lithic industries. Another site with evidence of very
early symbolic behavior (~45 ka) is Kara Bom,
which is located, like Denisova Cave, in the Russian
Altai Mountains (102).
What role did geographic and/or
paleoenvironmental variations play?
Modern humans dispersing out of Africa and
into Asia were able to adjust to a diversity of
new environments and often would have faced
a variety of geographic barriers (e.g., moun-
tain ranges, major riverways, deserts, and seas)
(30, 31, 34, 45, 46). Biological adaptations would
have facilitated dispersals during the Late Pleis-
tocene to high-altitude regions such as the Qinghai-
Tibetan Plateau by <30 ka (103) and far northern
Siberia by 32 ka (68, 71). Nevertheless, cultural
innovations would have been of even greater im-
portance than biological adaptations, simply be-
cause the latter may take generations to appear
in a population, whereas the former can appear
and spread quickly in a single generation. The
early development of footwear, as indirectly evi-
denced at the 40-ka Tianyuan Cave site in northern
China (104), is a case in point; this cultural trait
would have immediately facilitated human disper-
sals into colder environments (both latitudinally
and altitudinally).
Major natural events such as the Toba volcanic
super-eruption at 74 ka in Sumatra, Indonesia,
are surmised to have contributed to global cool-
ing, substantial landscape resculpting, and the
die-off of floras and faunas, leading to mam-
malian bottlenecks and extinctions (105, 106).
Ambrose (107) proposed that the Toba super-
eruption led to the extinction of archaic humans
and a population crash in modern H. sapiens.
However, on-the-ground research indicated that
terrestrial environments were reshaped more
subtly in India (108) and Africa (109) and that
Middle Paleolithic populations in the Indian sub-
continent survived the super-eruption (33, 108).
The present-day submergence of coastlines
containing traces of hominin occupation will hin-
der reconstruction of dispersal routes, partic-
ularly during major glacial periods when sea
levels were ~100 m below their current levels (54).
A good example of this is the eastern China
seaboard: During glacial periods, the coastal plain
would have extended outward 400 to 600 km and
connected areas such as the Shandong Peninsula
in eastern China with the Korean Peninsula, fa-
cilitating movement of a variety of animals, in-
cluding humans, across the dry Yellow Sea (94).
In the Korea-China case, there is good reason to
assume that a number of archaeological sites and
Pleistocene faunas are present in this submerged
former coastal plain. It is not clear that the same
argument can be made for all coastal regions in
Asia. For instance, proponents of the southern
coastal dispersal model argue that the reason that
there is a paucity of evidence of coastal occupation
is that the sites are now submerged (17, 55). How-
ever, the coastline along the Indian Ocean rim and
the western side of the Sunda Shelf has areas of
relatively steep drop-off, so that it is unlikely that
many coastal sites would be now submerged; more-
over, archaeological surveys in these steep coastal
shelves and in some uplifted shoreline areas have
failed to find Late Pleistocene coastal sites (31).
So what happened then?
Growing evidence indicates that modern human
dispersals out of Africa into Asia occurred by 60 ka
and afterward (Fig. 1B). Such dispersal events
across Eurasia are supported by a diversity of
studies from genetics and archaeology (11, 20, 28).
in modern human populations (43).
A later, major OoA event most likely occurred
some time around 60 ka and thereafter. This
later dispersal by larger and more demograph-
ically successful human populations masked
genetic traces of the earlier dispersals (73). These
dispersals across Asia occurred in both north-
erly and southerly directions (28). In the move
north, modern human foragers skirted the Qinghai-
Tibetan Plateau to its south to eventually reach
Siberia (70). At around the same time or soon after,
populations reached Europe. These northward-
traveling human populations carried an advanced
toolkit comprising microliths, blades, composite
tools, and symbolic objects (beads and colorants),
which eventually facilitated their dispersal and
successful establishment in higher latitudes and
altitudes. These foraging groups carrying a spe-
cialized microblade toolkit eventually made their
way to the Americas through Beringia (69). The
southern route includes possible movement path-
ways through the Indian subcontinent, mainland
Southeast Asia, and eventually as far north as
central China and southward into the Southeast
Asian islands and on to Australasia. Questions
remain as to why modern human foragers arriv-
ing in Southeast Asia discarded blade and micro-
blade stone tool industries, although evidence
indicates that communities had ground stone
technologies, which represented adaptations to
new environments (63). Further, there are ques-
tions concerning what happened when different
foraging groups, originating in areas to the north
and south, met in central China (34, 89).
Moving forward: Future directions of
Late Pleistocene Asian
paleoanthropology
A rigid definition of the OoA model positing that
modern humans dispersed from Africa only after
60 ka and simply replaced all indigenous popula-
tions (e.g., mid-Pleistocene Homo, Neandertals,
Denisovans, and H. floresiensis) with no inter-
breeding can no longer be considered valid. What
is needed now is to develop more detailed

Bae et al., Science 358, eaai9067 (2017) 8 December 2017 5 of 7

R ES E A RC H | R E V IE W
models that include the growing evidence for
early dispersals, contractions, extirpations, and
extinctions of human groups and lineages, while
recognizing that not all pre–60 ka dispersals
were evolutionary dead ends. Further, there are
still large swathes of territory in Asia that remain
largely unexplored, and previously identified sites
and materials are in need of renewed study. Fortu-
nately, the increasing number of multidisciplinary
research programs launched in Asia over the past
few decades has resulted not only in the reporting
of many important sites and findings, but also the
accumulation of information that fills in gaps in
the evolutionary record, thereby facilitating broader
interregional comparisons.
As attested to by the great interest generated
by the Central Asiatic Expeditions in the early
20th century, Asia was once considered to be
the cradle of mankind (14). Although important
finds from Europe and Africa over the course of
the past century have diverted attention, Asia is a
continent that has much to offer to research on
modern human origins, thereby shaping what we
think we know about our evolution and history.
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AC KNOWLED GME NTS
We thank the participants of the Wenner-Gren Symposium “Human
Colonization of Asia in the Late Pleistocene” for stimulating and
lively discussions in Sintra, Portugal. We acknowledge the
support over the years from the Wenner-Gren Foundation for
Anthropological Research, the National Geographic Society, the
U.S. National Science Foundation, the Academy of Korean Studies,
the College of Social Sciences and School of Pacific and Asian
Studies at the University of Hawaiʻi at Manoa (C.J.B.), the Max
Planck Institute for the Science of Human History (K.D. and M.D.P,)
and the European Research Council [ERC-2012-AdG-324139
PALAEOCHRON and ERC-2016-STG-715069 FINDER (K.D.) and
ERC-2011-AdG-295719 PALAEODESERTS (M.D.P.)]. We thank
L. Aiello and the anonymous reviewers for their comments and
recommendations.
10.1126/science.aai9067
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On the origin of modern humans: Asian perspectives
Christopher J. Bae, Katerina Douka and Michael D. Petraglia

Science 358 (6368), eaai9067.

DOI: 10.1126/science.aai9067

The peopling of Asia

In recent years, there has been increasing focus on the paleoanthropology of Asia, particularly the migration
patterns of early modern humans as they spread out of Africa. Bae et al. review the current state of the Late Pleistocene
Asian human evolutionary record from archaeology, hominin paleontology, geochronology, genetics, and
paleoclimatology. They evaluate single versus multiple dispersal models and southern versus the northern dispersal
routes across the Asian continent. They also review behavioral and environmental variability and how these may have

Downloaded from http://science.sciencemag.org/ on April 2, 2020

affected modern human dispersals and interactions with indigenous populations.
Science, this issue p. eaai9067

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