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The Late Pleistocene Colonization of South America: An Interdisciplinary

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 Review doi: 10.1111/j.1469-1809.2009.00537.x

The Late Pleistocene Colonization of South America: An

Interdisciplinary Perspective
Francisco Rothhammer1,2∗ and Tom D. Dillehay3,4
1
Instituto de Alta Investigación, Universidad de Tarapacá, Arica Chile
2
Programa de Genetica Humana, Instituto de Ciencias Biomédicas, Facultad de Medicina, Universidad de Chile, Santiago, Chile
3
Department of Anthropology, Vanderbilt University, Nashville, TN, USA
4
Instituto de Ciencias Sociales, Universidad Austral de Chile, Chile

Summary
In this article, we briefly review scenarios for the first peopling of the New World, including the Clovis First/Single
Origin, Tripartite, and Dual Migration models. We then discuss bioanthropological, molecular genetic, archaeological
and biogeographic evidence for the peopling of the continent. To conclude, we draw on different lines of evidence to
make inferences concerning the timing, direction, and type of early human dispersion in South America.

Keywords: Colonization of South America, multidisciplinary approach, provisional peopling model

Introduction The Clovis First/Single Origin Model

This essay considers the first human colonization of South
America from an interdisciplinary perspective, including
bioanthropological, genetic, archaeological and biogeograph-
ical evidence. Our objectives are twofold. First, we briefly
review models that have been proposed for the peopling of
the New World, with special reference to the southern hemi-
sphere, including the Clovis First/Single Origin (Hrdlička,
1937; Haynes, 2002), Tripartite (Greenberg et al., 1986), and
Dual Migration (Neves & Hubbe, 2005) models. Second, we
review some key results from bioanthropology, genetics, and
archaeology and their implications particularly regarding the
peopling of South America. Models based primarily on data
from a single discipline and/or methodological approach are
perhaps less likely to advance our understanding of initial hu-
man migrations. Our intention here is to build on previous
integrated analyses (e.g., Neves et al., 2003; Dillehay, 2008;
Fagundes et al., 2008; Goebel et al., 2008; González-José
et al., 2008) and to provide a broad critique and a simplifying
perspective.
∗ Origin model was later modified to fit the newly devised
Corresponding author: Prof. Francisco Rothhammer, Instituto de
Alta Investigacion, Universidad de Tarapaca, Arica 1001236, Chile.
Tel: 56-058-230334; Fax: 56-058-255371; E-mail: frothham@
med.uchile.cl
For decades the prevailing view among scholars interested
in the peopling of the Americas has been that Paleoindi-
ans arrived in the New World via Beringia from north-
east Asia during Clovis times around 11,500 years ago, and
that they were the direct ancestors of present-day Amerinds
(Dillehay, 1999; Dixon, 2001; Adovasio & Page, 2002;
Haynes, 2002; Meltzer, 2004; Goebel et al., 2008). This per-
spective of the first peopling of the Americas is termed the
Clovis First/Single Origin hypothesis, which was first pro-
posed by Ales Hrdlička (1937), a major participant in the
typology and race debates in biological anthropology during
the early part of the 20th century (Blakey, 1987).
After the discovery of “Folsom Man” in the 1920s, Hrdlicka
vigorously refuted all claims for any human presence more
than 4000 years old in the New World. However, the dis-
covery of the Clovis site in 1932 turned him into one of the
leading advocates in revising the perception of the first hu-
mans entering the New World and he conceded that they
may have arrived in late Pleistocene times. When Clovis
projectile points later appeared at numerous sites in North
America, often associated with the bone remains of extinct
mammals such as mammoth (dated by radiocarbon means to
ca. 11,000 years ago) the idea of Clovis Paleoindians having
been the first Americans was born. The Clovis First/Single
chronological and cultural schemes for the initial peopling of
the Americas (e.g., Haynes, 2002). More recently, archaeo-
logical evidence from various sites scattered throughout the

540 Annals of Human Genetics (2009) 73,540–549 

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Americas and dating from before Clovis times (e.g.,
Dillehay, 1997; McAvoy & McAvoy, 1997; Adovasio et al.,
1999; cf. Goebel et al., 2008), has overturned the “Clovis-
First” paradigm, establishing that the first humans arrived in
North and South America in pre-Clovis times (c.f., Bryan,
1986; Dillehay, 1997, 2000; Adovasio & Page, 2002; Stanford
& Bradley, 2002; Meltzer, 2004, 2006; Goebel et al., 2008).
Shortly after the discovery of the Clovis site, the question
arose as to how early hunters had reached the mid-latitudes
of North America during the Wisconsin Glaciation (between
25,000 and 10,000 years B.P.), at a time when the north-
ern half of the continent was thought to be buried beneath
two extensive ice sheets called Laurentide and Cordilleran.
In 1933, W.A. Johnston proposed the existence of an an-
cient passage between the two glaciers, later termed the “ice-
free corridor” by Antevs, which stretched from the Yucon
to northern Montana (Jackson et al., 1997). New geological
and archaeological information has doubted the existence of
this “corridor” (Jackson et al., 1997; Mandryk et al., 2001;
Clague et al., 2004), however, leading researchers to con-
sider other routes of entry from Beringia. Several investigators
(Fladmark, 1979, 1983; Dixon, 1993; Gruhn, 1994; Erland-
son, 2002) have suggested that the earliest human migration to
the Americas may have occurred along the north-west coast
through the use of watercraft to navigate the oceanic edges
of the continent. Coastal stretches may have been deglaciated
by about 16,000 B.P. (Mann & Peteet, 1994; Mandryk et al.,
2001). The coastal entry hypothesis has recently gained in-
creased support particularly as a consequence of (1) geological
research in Alaska and the Yukon (e.g., Jackson et al., 1997;
Mandryk et al., 2001), which has established new chronolo-
gies for major climatic events in the region and further ques-
tioned the existence of the corridor model, (2) the more
general acceptance of a pre-Clovis culture, and (3) the dis-
covery of several early coastal sites along the north-west coast
of North America and the Pacific coast of South America
(Keefer et al., 1998; Sandweiss et al., 1998; Dixon, 2006:
Dillehay et al., 2008).
The Tripartite Model
In the 1980s, Greenberg et al. (1986) and Turner (1987) em-
ployed linguistic, dental, and genetic data to suggest that the
Americas were peopled by three different migrations: (1) the
early Amerindians who first reached the interior of Alaska and
the Yukon and then spread to some territories of Canada and
eventually to the rest of the New World; (2) subsequently, the
north-west Pacific coast was colonized by Na-Dene speakers;
and (3) lastly, the Arctic was occupied by Eskimos. Although
this hypothesis received considerable attention in attempting
to explain the peopling of the Americas from an interdisci-
plinary perspective, it was later strongly challenged on linguis-

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Late Pleistocene Colonization of South America
tic, bioanthropological and genetic grounds (Morell, 1990;
Neves & Pucciarelli, 1991; Szathmáry, 1993a,b; Lahr, 1995;
Merriwether et al., 1995) and never had a widespread and
lasting impact on the way researchers considered the peopling
of South America.
The Dual Migration Hypothesis
During the 1990s, North American patterns of cranial vari-
ability were examined and found to exhibit heterogeneous
morphological trends, supporting the view that different pop-
ulations entered the New World at various time intervals. Fur-
thermore, although the closest morphological affinities were
with Asian populations, the data could not be used to spec-
ify the time of entry or the number of founding populations
(Steele & Powell, 1992; Jantz & Owsley, 2001). Recently, a
group of South American researchers (Neves & Pucciarelli,
1990, 1991; Neves et al., 2003; Neves & Hubbe, 2005) have
suggested, on the basis of a comparison of skulls from Lagoa
Santa, Brazil, with late prehistoric and modern Amerindian
crania, that craniometric variation in the New World re-
veals two different, discrete patterns, termed Paleoamerican
and Amerindian. Whereas Paleoamericans tend to morpho-
logically resemble present-day Australians, Melanesians, and
Sub-Saharan Africans, late prehistoric and modern Amerinds
show a greater resemblance to northern Asians. The propo-
nents of this hypothesis claimed that two successive migra-
tions, stemming from different geographical and chronologi-
cal sources, were responsible for the observed patterns of vari-
ation. According to this model, similarities among Australians,
Late Pleistocene Asians (e.g., the Zhoukoudien Upper Cave,
Minatogawa and Liujiang skulls) and Paleoamericans can be
explained by descent from South-eastern Asia (Neves et al.,
2003). Paleoamericans are believed to have arrived following
a terrestrial route by 15,000 B.P., whereas Amerindians came
about 4000 years later along a similar route (Hubbe et al.,
2003; Pucciarelli, 2004).
Since the Lagoa Santa skulls have gained a protagonist po-
sition in South American physical anthropology, it is useful to
review their history in more detail. The crania were initially
collected by the Danish explorer Peter Lund in the caves of
Lagoa Santa, Brazil, between 1835 and 1844. Lund, based
on the Paleolithic chronology of Europe, assigned a great an-
tiquity to the skulls; this was later challenged by McCown
(1963) and others. Subsequently, inspired by the putative an-
tiquity of the long and narrow headed Lagoa Santa crania, the
French anthropologist Paul Rivet published the morphome-
trical analysis of 17 skulls of the same “type” selected from
a series of 78 undeformed specimens of indeterminate age
collected from several rock shelters in Paltacalo, southern
Ecuador (McCown, 1963). Shortly thereafter, Rivet pub-
lished a revision of the “racial” position of a series of modern
Annals of Human Genetics (2009) 73,540–549 541
 F. Rothhammer and T. D. Dillehay
Pericú skulls from the Baja California Peninsula (Rivet, 1909).
The Baja skulls were suggested to belong, together with the
Paltacalo skulls, to different branches of a Lagoa Santa “race”,
and to resemble Australian and Melanesian crania. (We note,
parenthetically, that the same modern Pericú skulls were re-
cently used as evidence for the existence of a “Paleoamerican
pocket” in the lower part of Baja California, which apparently
survived in the area due to hypothetical climatic changes and
geographic isolation during the middle Holocene (González-
José et al., 2003).) The Paltacalo skulls eventually acquired a
“fuzzy aura of antiquity” among scholars and became, along
with the Lagoa Santa crania, the homotypes of the long and
narrow-headed ancient Lagoa Santa or Paleoamerican “race”
(c.f., McCown, 1963). McCown described in detail further
developments of what he called “a curious intellectual edi-
fice”, and concluded that “the initial demonstration of Rivet’s
race of Lagoa Santa is based upon material that a properly
trained modern anthropologist cannot but regard as utterly
inadequate” (McCown, 1963).
In order to cope with the problems derived from the dis-
puted antiquity of the Lagoa Santa skulls, for which only two
radiocarbon dates have been reported (9720 ± 128 and 9028
± 120 B.P. non-calibrated; Neves & Hubbe, 2005), Neves
generated a more reliable chronological framework of the
skulls in the 1990s. Two skulls of the studied samples now
date back to ca. 11,000 B.P., namely the Confins skull from
Lapa Mortuária (cave) (Walter et al., 1937) and “Luzia” from
Lapa Vermelha IV (Prous & Fogaça, 1999). Both skulls were
reported to have been dated on charcoal, associated either
stratigraphically or directly with the burials (Neves & Hubbe,
2005). Since the first studies were based on a very small sam-
ple of crania, additional efforts to increase sample size were
subsequently made, assembling a collection of 81 specimens
from Central Brazil. The vast majority of the skulls (n = 74)
were found to date between 8000 and 8500 years B.P. (Neves
& Hubbe, 2005). An exploratory statistical comparison with
Howells’ database (Howells, 1996) established that the Lagoa
Santa skulls resemble present-day Australian/ Melanesian and
Africans confirming the results reported at the beginning of
the last century by Rivet for his Lagoa Santa “race”. Rivet
(1943) interpreted this similarity as proof of an ancient migra-
tion to America across the Pacific Ocean, whereas the pro-
ponents of the dual migrational hypothesis have postulated a
terrestrial route from Africa to America via Asia (Neves &
Hubbe, 2005).
In order to be phylogenetically informative, cranial varia-
tion should be selectively neutral and not strongly affected by
stochastic microevolutionary forces (drift). A number of re-
cent studies indicate that in modern human populations skull
size and shape are not predominantly the result of selection,
but mainly result from the interaction between gene flow
and drift and, to a lesser extent, mutation (Relethford, 2002;
Ackermann & Cheverud, 2004; González-José et al., 2008).
542 Annals of Human Genetics (2009) 73,540–549
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Operating under the assumptions of a migration/founder
model, Powell & Neves (1999) examined a series of Pale-
oindian and Archaic crania from North and South America
and compared these data to a large world-wide sample of
late Holocene remains to assess within- and among-group
variation and to determine how the observed pattern could
be interpreted in terms of population history and structure.
Powell & Neves (1999) suggested that their cranial data were
consistent with the first Americans having been derived from
an undifferentiated Asian population that was not ancestral
to modern American Indians. No doubt, this result can be
accommodated to support multiple migration scenarios. Nev-
ertheless, according to the same authors, the assumptions un-
derlying the model were not very realistic and when the
data were analyzed controlling for the effects of drift, the
first Americans were no longer differentiated from modern
Amerindians.
Classical Genetic Data
Turning to information provided by synthetic gene-
geography graphs based on blood group and protein data,
we note that maps of South America based on a small num-
ber of polymorphic loci are, as expected, not very informative
(O’Rourke & Suarez, 1985). Increasing the number of loci
results in maps where the first principal component (roughly
50% of gene frequency variation) reveals clines extending
from north-western to south-eastern South America that can
be interpreted as indicative of very early population displace-
ments from the Isthmus of Panama to the Brazilian coast and
down to the southern cone of the continent (Rothhammer
& Silva, 1992; Rothhammer et al., 1997). Maps based on the
second and third principal component exhibit clines which
are more difficult to interpret, but which may indicate pop-
ulation movements during the Late Archaic/Early Formative
along available waterways in search of new alluvial land in the
Amazon flood plain, as claimed by Lathrap (1970). Cavalli-
Sforza et al. (1994), suggested an early southward migration
along the western side of the Andes, consistent with the inter-
pretation that modern speakers of Andean languages may rep-
resent descendants of the first occupiers of the region. Lastly,
an analysis of 31 protein systems obtained in 29 Amazonian
populations agreed with a previously observed differentiation
between northern and southern Brazil, suggesting that the
Amazon river could have constituted a barrier to north-south
gene flow or that genetically differentiated groups may have
entered the region from the west (Callegari-Jacques et al.,
1994)
Molecular Genetic Evidence
Mitochondrial and Y-chromosome variation in North and
South America indicate unambiguously that all Native

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Americans came from Asia and not from Melanesia, Australia,
Africa, or Europe (Karafet et al., 2006; Merriwether, 2006;
Wang et al., 2007). Five mtDNA (A; B; C; D and X) and
two Y-chromosome (C and Q) founding haplogroups, which
are all present among native populations of Siberia account
for the molecular variability observed among extant Native
American populations (Derenko et al., 2001; Zegura et al.,
2004; Starikovskaya et al., 2005). The complete sequencing
of 84 mtDNAs have filled gaps in the internal sequence vari-
ation of A,C and D, contributing to improve the resolution of
the mtDNA phylogeny in Siberia-Beringia (Volodko et al.,
2008) Haplogroup X includes many lineages, some of which
are present in Europe and Asia. Nevertheless, the lineage
found in American Indians is different from those observed
in Eurasia (Merriwether, 2006). Ancient mtDNA analysis has
until now yielded the same founding haplogroups described
in extant populations, thus confirming the genetic continu-
ity between extinct and extant Native American populations
(Stone & Stoneking, 1998; Kemp et al., 2007). Consequently,
many migration models based on the presence or absence of
certain haplogroups have lost much explanatory usefulness
(c.f., Goebel et al., 2008).
In an extensive study on patterns of genetic variation in Na-
tive Americans from both North and South America, Wang
et al. (2007) reported the results of the analysis of 678 mi-
crosatellite markers genotyped in 422 individuals represent-
ing 24 Native American populations. The analysis of the data
revealed a progressive reduction of variability starting from
Africa through Asia and down from the Bering Strait to North
and South America, consistent with a serial founding African-
origin model of human evolution (Prugnolle et al., 2005;
Ramachandran et al., 2005). Noteworthy was the observa-
tion in South America of a west-to-east difference in genetic
diversity, showing that eastern (Brazilian) populations had the
lowest levels of heterozygosity. This pattern was also observed
with mitochondrial DNA (Fuselli et al., 2003; Lewis et al.,
2004) and Y-chromosome markers (Tarazona-Santos et al.,
2001). The fact that Brazil exhibits the lowest levels of varia-
tion suggests an initial colonization of western South America
and a subsequent peopling of the eastern part by western sub-
groups, confirming the interpretation of the results of a pre-
vious genetic analysis of Amazonian tribes (Callegari-Jacques
et al., 1994).
Colonization routes also were examined by Wang et al.
(2007), by evaluating the relationship between heterozygosity
and “effective” geographic distances. When coastlines were
treated as preferred routes, there was an increase in nega-
tive correlation between heterozygosity and distance from
the Bering Strait. A relative genetic similarity between An-
dean and Mesoamerican populations was also observed by
Wang et al. (2007) and was interpreted as consistent with
an early coastal colonization. An alternative interpretation
is that the two most developed pre-Columbian cultures in

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Late Pleistocene Colonization of South America
the Americas flourished precisely in Mexico and Peru, and
that the observed genetic affinities could be the result of late
pre-Hispanic (or even early Hispanic) contact between them.
We note in passing, that anecdotal information reported by
the Spanish chroniclers suggests that in Peru, under the rule
of Tupac Inca Yupanqui, long transpacific voyages along the
Pacific coast and out to sea were organized using large bal-
sas (rafts) (Rivet, 1943; Cabello-Valboa, 1951; Pietschmann,
1906).
Archaeological Data and Biogeography
of South America
The late Pleistocene archaeological record of South Amer-
ica differs from that of North America in many ways
(Bryan, 1973, 1986; Dillehay, 1999; Miotti et al., 2003;
Borrero, 2006). Among these, there is an absence in South
America of a continent-wide stone tool style like Clovis and
of evidence for the long-distance movement of raw lithic
material. The presence of sharply contrasting environments
in the southern hemisphere favoured the establishment of
distinct regional cultural traditions at the outset of human
dispersion. Furthermore, the extensive ice sheets covering
high latitudes limited the movement of colonizers in North
America in contrast with South America where the glacial
effect was confined to patchy high altitude and latitude areas
(Clapperton, 1993). The archaeological evidence of human
entry into South America before 15,000 B.P. is weak and
only presumed at this time. However, given the evidence
confirming the presence of humans probably centuries before
12,500 B.P., the likely entry date should lie between 15,000
and 13,500 B.P.
Between at least 11,000 and 10,000 B.P. South America
witnessed several major changes, including the increased use
of coastal resources, demographic concentration in major river
basins, and modification of landscapes and plant and animal
distributions. Some 1000 years later an increase of site den-
sity, technological diversification, plant domestication, and
appearance of ritual practices indicate that the initial impulses
toward cultural complexity had developed in South America,
probably within only a few millennia after the initial arrival of
colonizers (Dillehay, 2000; Lavallee, 2000). No doubt, sharp
contrasting environments characterized the continent dur-
ing the late Pleistocene and Pleistocene to Holocene tran-
sition and contributed to cultural developments that show a
steady shift away from sub-continental uniformity and toward
the establishment of distinct regional, cultural, and especially
technological traditions (Bryan, 1973, 1986; Prieto, 1996;
Dillehay, 1999; Borrero, 2006). The impact of climatic and
environmental changes on these developments is not well un-
derstood. There is evidence of a significant temperature rise
between 15,000 and 14,000 B.P. and a subsequent warm-
Annals of Human Genetics (2009) 73,540–549 543
 F. Rothhammer and T. D. Dillehay
ing trend lasting until 10,500 B.P. (Rull, 1996; Prieto, 1996;
Latrubesse & Rambonell, 1994; Weng et al., 2004; Bush &
John, 2006).
South America is biogeographically divisible into four ma-
jor areas: the Andes, the tropical and temperate plains of
Colombia, Venezuela, and Brazil, the highlands of east Brazil
and the Guyanas, and the southern Pampa and Patagonian
grasslands. These divisions are important for considering hy-
pothetical migration routes into the southern hemisphere
(c.f., Bennett & Bird, 1964). The Andean mountain chain,
the vast Amazonian basin, and the southern grasslands were
doubtless a combined barrier for isolating some human pop-
ulations geographically and genetically, regularly disrupting
gene flow between geographically contiguous populations,
and producing some of the regional skeletal and genetic dif-
ferences discussed earlier. Although foraging mobility un-
doubtedly continued throughout the terminal Pleistocene to
early Holocene periods, reduced mobility in some regions
probably derived from the increasing use of local resources
constitutes one potential explanation for the bioanthropolog-
ical heterogeneity observed across the continent. Both ge-
netic and morphological signals of marked diversity between
the Andes and the eastern lowlands of the continent, gener-
ally agree with differences observed in the technological and
subsistence patterns in these two regions.
Continental ice sheets started to melt and the sea levels
began to rise between 14,000 and 13,000 B.P. The Pacific
and Atlantic shelves and many areas in Tierra del Fuego were
flooded, as were any archaeological sites, making the study of
early migration routes along the coasts very difficult. These
and other late Pleistocene environmental changes introduced
significant resource-related changes that likely favoured social
learning and bonding and technological change, as evidenced
by a wide array of local and regional stone tool technolo-
gies and subsistence patterns in place by at least 10,500 years
ago (Kipnis, 1998; Dillehay, 2000; Salemme & Miotti, 2002;
Flegenheimer et al., 2006; León-Canales, 2007). The broad
network of environmental responses by people across the con-
tinent clearly included more than the development of social
bonding and strategies for sharing resources. There is so much
diversity across the continent at this time and later that it is
difficult to say to what degree the eastern lowlands were
culturally different from the Andean region or the southern
grasslands were from the Amazon basin or northern tropical
forests. Regional technological diversity suggests that there
were specific responses by different groups to the great vari-
ations in local habitats and raw material availability that ex-
isted across the continent, especially in the Andean region.
Technology offered a flexible, rapid means of responding to
climate and other changes at a range of scales from daily
needs of individuals to long-term trends experienced over
generations.
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Several sub-continental studies have emphasized the insta-
bility of climates during terminal glacial, deglaciation, and
the transition to the early Holocene from ca. 13,000 to
10,000 years ago (Bennett et al., 2000; Bush & John, 2006;
Lovell & Kelly, 2008). For humans, these cyclical periods of
climatic stability and instability would have placed additional
selective pressures on integrated biological and behavioural
responses to increased resource variability. At the continen-
tal scale, populations distributed across Amazonia or along the
Pacific coastal shelf during warm deglaciation, possibly would
have become geographically isolated during times of aug-
mented aridity. Increased precipitation and vegetation growth
in some areas after 11,000 B.P. may have pushed some pop-
ulations outward towards the better vegetated and watered
areas, such as the Amazonian rainforest and the mid-latitudes
and western altitudes of the continent. Such repeated cycles
and disruptions of landscapes were doubtless a force for isolat-
ing populations geographically, biologically, and consequently
genetically, regularly disrupting gene flow even between con-
tiguous populations and more so between those on the east
and west sides of the continent. At sub-continental scales,
expansions and contractions of environments during the ter-
minal Pleistocene and early Holocene period (ca. 12,000–
10,000 B.P.) would have drawn in or pushed out human pop-
ulations from north to south, or vice versa, as well as eastwards
into the Amazon basin during times of expansion or retreat
from aridity, for example. The disruption of contact between
regional populations combined with occasional severe down-
turns, causing local animal and possibly human extinctions,
also would have contributed to the development of regional
differences in skeletal morphology, gene pools, and social and
technological behaviours. Although social processes must also
be taken into consideration to explain these differences in the
morphological, genetic, and archaeological records, for which
there is currently no hard evidence, the periodicity and ex-
tremes of late Pleistocene climatic cycles must have underlain
a process of accretionary or mosaic emergence of human and
cultural variation within South America.
Considering the various sources of evidence discussed
above one can outline a possible scheme of human dispersion
in South America. Hunters and gatherers who migrated to
South America via the Isthmus of Panama could have entered
the Andean highlands by way of the Cauca and Magdalena
river valleys which flow from south to north in Colombia.
Once adjusted to the highland environment there would have
been few barriers for moving further south, although some
high altitude areas were covered by glaciers. Some groups may
also have migrated eastward following the Caribbean rim of
Venezuela, the Guyanas, and north-east Brazil and others into
the interior of Venezuela and afterwards south-east or south-
west along several large river systems into the Amazon basin.
People may also have migrated along the Pacific coast down to

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 Late Pleistocene Colonization of South America

Chile, following favourable fishing localities and using water-
craft (Rothhammer & Silva, 1992; Cavalli-Sforza et al., 1994;
Keefer et al., 1998; Sandweiss et al., 1998; Stothert, 1998;
Wiesner, 1999; Wang et al., 2007). From the Amazon basin
and/or the Andes of north-west Argentina, people could have
entered the open parkland country of eastern Brazil and also
could have spread throughout the Pampas and Patagonia. As
mentioned above, genetic data from Native Brazilian popula-
tions suggests that the Amazon River constituted a barrier to
north-south gene flow. There is also the possibility that ge-
netically divergent groups may have entered the region from
Colombia, Venezuela, and the Guyanas in the north and from
Argentina in the south (Fig. 1).
There is evidence in parts of north-western and central
South America that several plant species were deliberately ma-
nipulated and possibly domesticated in terminal Pleistocene
times (Piperno & Pearsall, 1998; Piperno & Stothert, 2003).
What is not understood is whether there are any connections
between broad-spectrum diets in environments where poten-
tially domesticable plants occurred, specific resource zones,
and social changes that might have favoured the first impulses
toward food production. Similar connections need to be as-
certained for the kinds of human and camelid interactions
that took place during this period in the puna and altiplano
zones of the central and south-central Andes. Diets, resource
zones, and local human population histories constitute only

Figure 1 Hypothesized migration routes into and throughout South America, as well as
some of the major regional archaeological sites dating to the late Pleistocene period.


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 F. Rothhammer and T. D. Dillehay
some of the variables required to understand the development
of early cultural diversity, food production within a relatively
short period of time after humans arrived, in comparison to
the Old World, and how ecological constraints and opportu-
nities played out in setting the stage for subsequent sedentary
and later more complex societies in South America.
Conclusion
South America was probably colonized between at least
15,000 and 13,500 B.P. most likely by one migration wave.
Archaeologists have noted that the peopling of the south-
ern hemisphere differed in many ways from that of North
America. Most significant has been the observation that no
single culture dominated the South American continent the
way Clovis did. Several factors likely account for this early
diversity including geographic barriers to human movement
(Andean Highlands and Amazon River), climatic changes,
shifts in resource areas, new settlement patterns, new lifestyles,
innovative technologies (e.g. domestication of plants), and
new social organizations, among others. In addition, the rel-
ative isolation derived from an absence of later migrants, as
seems to have occurred for North America, may have con-
tributed to early biological diversification and cultural com-
plexity of South America.
The west and east sides of South America have different
archaeological records, different chronologies and different
human histories. No doubt, these factors are reflected in dis-
tinctive patterns of genetic, morphological and cultural vari-
ation. Although South America still does not have sufficient
interdisciplinary data from archaeology, genetics, bioanthro-
pology and biogeography to precisely model the timing and
processes involved in the first peopling of the continent, a ten-
tative scheme of dispersal of first colonizers can be outlined
(Fig. 1).
The conclusion which may be drawn from this review of
various interdisciplinary approaches to understand the initial
peopling of South America is a positive one, with contin-
ued new insights being made in analytical techniques, yield-
ing more reliable results over a greater temporal and spacial
framework. Whereas any one approach may produce new
and different data and insights, as well as misleading results,
and various degrees of variation are likely among the findings
they provide, new data and techniques should continue to
reveal patterns not only specific to South America, but oth-
ers that relate to hemisphere-wide processes of migration and
colonization. In fact, theoretical generalizations of studies of
the first Americans have been scanty and their contribution
to global issues discrete. In this context, we wish to empha-
size the uniqueness of the colonization of South America as a
model to contribute to the understanding of the more general
process of adaptive radiation of populations anywhere.
546 Annals of Human Genetics (2009) 73,540–549
Journal compilation 
Acknowledgements
We thank the Editor and anonymous reviewers for helpful advice.
We also acknowledge gratefully the collaboration of Will Wilcox
in the elaboration of the image showing location of archaeo-
logical sites and possible migration routes. Financial support was
provided by Fondecyt 1095006 and Convenio de Desempeño
UTA/Mecesup-2 grants.
References
Ackermann, R. R. & Cheverud, J. M. (2004) Detecting genetic
drift versus selection in human evolution. Proc Natl Acad Sci USA
101, 17946–17951.
Adovasio, J. M., Pedler, D., Doanhue, J. & Stuckenrath, R. (1999)
No Vestige of a Beginning nor Prospect for an End: Two Decades
of Debate on Meadowcroft Shelter. In: Ice Age Peoples of North
America (eds. R. Bonnichsen and K. Turmire), pp. 416–431. Ore-
gon State University Press.
Adovasio, J. & Page, J. (2002) The First Americans: In Pursuit of Ar-
chaeology’s Greatest Mystery. New York: Random House.
Bennett, W. C. & Bird, J. (1964) Andean Culture History. Garden
City, New York: The Natural History Press.
Bennett, K. D., Haberle, S. G. & Lumley, S. H (2000). The Last
Glacial-Holocene Transition in Southern Chile. Science 290, 325–
328.
Blakey, M. L. (1987) Intrinsic social and political biology in the his-
tory of American Physical Anthropology. Critique of Anthropology
7, 7–35.
Borrero, L. (2006) Paleoindians without Mammoths and Archae-
ologists without Projectile Points? The Archaeology of the First
Inhabitants of the Americas. In: Paleoindian Archaeology: A Hemi-
spheric Perspective (eds. J. E. Morrow & C. Gnecco), pp. 9–20.
University Press of Florida.
Bryan, A. (1973) Paleoenvironments and cultural diversity in Late
Pleistocene South America. Q Res. 3, 237–256.
Bryan, A. (1986) Paleoamerican prehistory as seen from South
America. In: New Evidence for the Pleistocene Peopling of the New
World (ed. A. Bryan), pp. 1–14. Orono, ME: Center for the Study
of Early Man.
Bush, M. B. & John R. F. (2006) Tropical Rainforest Responses to
Climatic Change (eds. M. B. Bush & R. F. John). Berlin: Springer.
Cabello-Valboa, M. (1951) Miscelánea antártica. Una historia del antiguo
Perú, pp. 322–324. Lima.
Callegari-Jacques, S. M., Salzano, F. M., Weimer, T. A., Hutz,
M. H., Black, F. L., Santos, S. E., Guerreiro, J. F., Mestriner,
M. A. & Pandey, J. P. (1994) Further blood genetic studies on
Amazonian diversity-data from four Indian groups. Ann Hum Biol
21, 465–481.
Cavalli-Sforza, L. L., Menozzi, P. & Piazza, A. (1994) The history and
geography of human genes. Priceton: Princeton University Press.
Clague, J. J., Mathewes, R. W. & Ager, T. A. (2004) In: Entering
America: Northeast Asia and Beringia before the Last Glacial Maximum
(ed. D. B. Madsen), pp. 63–94. Salt Lake City: University of Utah
Press.
Clapperton, C. M. (1993) Quaternary Geology and Geomorphology of
South America. Amsterdam: Elvesier.
Derenko, M. V., Grzybowski, T., Malyarchuk, B. A., Czarny, J.,
Miścicka-Sliwka, D. & Zakharov, I. A. (2001) The Presence of
Mitochondrial Haplogroup X in Altaians from South Liberia. Am
J Hum Genet 69, 237–241.

C 2009 The Authors
C 2009 Blackwell Publishing Ltd/University College London

Dillehay, T. D. (1997) Monte Verde. A Late Pleistocene Settlement in
Chile. Vol. 2. The Archaeological Context and Interpretation, p. 1071.
Washington, DC, London: Smithsonian Institution Press.
Dillehay, T. D. (1999) The late Pleistocene Cultures of South Amer-
ica. Evol Anthropol 7, 206–216.
Dillehay, T. D. (2000) The Settlement of the Americas. New York: Basic
Books.
Dillehay, T. D. (2008) Probing Deeper into first American studies.
Proc Natl Acad Sci USA 106, 971–978.
Dillehay, T. D., Ramirez, C., Pino, M., Collins, M., Rossen, J.
& Pino-Navarro, D. (2008) Monte Verde: Seaweeds, food, and
medicine and the peopling of the Americas. Science 325, 1287–
89.
Dixon, E. J. (1993) Quest for the Origins of the First Americans. Albu-
querque: University of New Mexico Press.
Dixon, E. J. (2001) Human colonization of the Americas: Timing,
technology and process. Quatenary Sci Rev 20, 277–299.
Dixon, E. J. (2006) Peopling of the Americas. How and when did
people first come to North America? Athena Review 3, 2.
Erlandson, J. (2002). Anatomically modern humans, maritime voy-
aging, and the Pleistocene colonization of the Americas. In: The
First Americans, the Pleistocene Colonization of the New World (ed.
N. G. Jablonski), No. 27, pp. 59–92. Memoirs of the California
Academy of Sciences.
Fagundes, N. J., Kanitz, R., Eckert, R., Valls, A. C., Bogo, M.
R., Salzano, F. M., Smith, D. G., Silva, W. A., Jr., Zago, M. A.,
Ribeiro-dos-Santos, A. K., Santos, S. E., Petzl-Erler, M. L. &
Bonatto, S. L. (2008) Mitochondrial population genomics sup-
ports a single pre-Clovis origin with a coastal route for the peo-
pling of the Americas. Am J Hum Genet 82, 583–92.
Fladmark, K. R. (1979) Routes: Alternative Migration Corridors
for Early Man in North America. American Antiquity 44, 55–69.
Fladmark, K. R. (1983) Times and Places: Environmental Corre-
lates of Mid-to-Late Wisconsin Human Population Expansion in
North America. In: Early Man in the New World (ed. R. Shutler),
pp. 13–42. Beverly Hills: Sage Publication.
Flegenheimer, N., Bayon, C. & Pupio A. (2006) Llegar a un Nuevo
Mundo: La Arqueologı́a de los Primeros Pobladores del Actual
Territorio Argentino. Museo y Archivo Histórico Municipal, Bahia
Blanca.
Fuselli, S., Tarrazona-Santos, E., Dupenloup I., Soto, A. & Luiselli,
D. (2003) Mitocondrial DNA diversity in South America and the
genetic history of Andean highlanders. Mol Biol Evol 20, 1682–
1691.
Goebel, T., Waters, M. & O’Rourke, R. (2008) The late pleistocene
dispersal of modern humans in the Americas. Science 319, 1497–
1502.
González-José, R., González-Martı́n, A., Hernández, M., Puccia-
relli, H. M., Sardi, M., Rosales, A. & Van Der Molen, S. (2003)
Craniometric evidence for Palaeoamerican survival in Baja Cali-
fornia. Nature 425, 62–65.
González-José, R., Bortolini, M. C., Santos, F. R. & Bonatto, S. L.
(2008) The peopling of America: craniofacial shape variation on
a continental scale and its interpretation from an interdisciplinary
view. Am J Phys Anthropol 137, 175–187.
Greenberg, J. H., Turner, C. G. & Zegura, S. L. (1986) The settle-
ment of the Americas: a comparison of the linguistic, dental and
genetic evidence. Curr Anthropol 27, 477–495.
Gruhn, R. (1994) The Pacific Coast Route of Initial Entry: An
Overview. In: Method and Theory for Investigating the Peopling of
the Americas (eds. R. Bonnichsen & D. G. Steele). Corvallis, OR,
USA: Center for the Study of the First Americans, Oregon State
University.

C 2009 The Authors
Journal compilation 
C 2009 Blackwell Publishing Ltd/University College London
Late Pleistocene Colonization of South America
Haynes, G. A. (2002) The early settlement of North America: The Clovis
Era. Cambridge: Cambridge University Press.
Howells, W. W. (1996) Howells’ craniometric data on the Internet.
Am J Phys Anthropol 101, 441–442.
Hrdlička, A. (1937) The origin and antiquity of the American In-
dian. Annu Rep Smithson Inst Washington 1923, 481–494.
Hubbe, M., Mazzuia, E. T. A., Atui, J. P. V. & Neves, W. A. (2003)
A primeira descoberta de America. Sao Paulo: Sociedade Brasileira de
Genética.
Jackson, L. E., Jr., Phillips, F. M., Shimamura, K. & Little, E. C.
(1997) Cosmogenic 36Cl dating of the Foothills erratics train,
Alberta, Canada. Geology 25, 195–198.
Jantz, R. L. & Owsley, D. W. (2001) Variation among early North
American crania. Am J Phys Anthropol 114, 146–155.
Karafet, T. M., Zegura, S. L. & Hammer, M. F. (2006) In: Handbook of
North American Indians Vol. 3: Environment, Origins and Populations
(eds. D. Ubelaker, W. C. Sturtevant & D. Stanford), pp. 831–839.
Washington, DC: Smithonian Institution Press.
Keefer, D. K., deFrance, S. D., Moseley, M. E., Richardson, J.
B., III, Satterlee, D. R. & Day-Lewis, A. (1998) Early maritime
Economy and El Niño events at Quebrada Tacaguay, Perú. Science
281, 1830–1832.
Kemp, B. M., Malhi, R. S., McDonough, J., Bolnick, D. A., Esh-
leman, J. A., Rickards, O., Martinez-Labarga, C., Johnson, J. R.,
Lorenz, J. G., Dixon, E. J., Fifield, T. E., Heaton, T. H., Worl, R.
& Smith, D. G. (2007) Genetic analysis of early holocene skeletal
remains from Alaska and its implications for the settlement of the
Americas. Am J Phys Anthropol 132, 605–621.
Kipnis, R. (1998) Early hunter-gatherers in Americas: Perpsectives
from central Brazil. Antiquity 72, 581–592.
Lahr, M. M. (1995). Patterns of modern human diversification: im-
plications for Amerindian origins. Am J Phys Anthropol 38, 163–
198.
Lathrap, D. W. (1970) The Upper Amazon. Southampton: Thames
and Hudson.
Latrubesse, E. M. & Rambonell, C. (1994) A climatic model for
southwestern Amazonia in late glacial times. Quaternary Int 21,
163–169.
Lavallee, D. (2000) The First South Americans. Salt Lake City: Uni-
versity of Utah Press.
León-Canales, E. (2007) Orı́genes Humanos en los Andes del Perú.
Lima: Universidad San Martin de Porres.
Lewis, C. M., Tito, R. Y., Lizarraga, B. & Stone, A. C. (2004) Land,
language and loci : mt DNA in Native Americans and the genetic
history of Peru. Am J Phys Anthropol 127, 351–360.
Lovell, T. V. & Kelly, M. A. (2008) Was the yonger dryas global?
Science 321, 348–349.
Mandryk, C. A. S., Josenhans, H., Fedje, D. W. & Mathewes, R.
W. (2001) Late Quaternary paleoenvironments of Northwestern
North America: implications for inland versus coastal migration
routes. Quaternary Sci Rev 20, 301–314.
Mann, D. H. & Peteet, D. M. (1994) Extent and timing of the
last glacial maximum in southwestern Alaska. Quaternary Res 42,
136–148.
McAvoy, J. M. & McAvoy, L. D. (1997) Archaeological Investi-
gations of Site 44SX202, Cactus Hill, Sussex County, Virginia.
Virginia Department of Historic Resources, Research Report 8,
Richmond, VA.
McCown, T. D. (1963) Handbook of South American Indians, Vol. 6,
pp. 1–9. New York: Cooper Square Publishers.
Meltzer, D. J. (2004) Peopling of North America. In: The Quaternary
period in the United States (eds. A. Gillespie, S. C. Porter & B.
Atwater), pp. 539–63. New York: Elsevier Science.
Annals of Human Genetics (2009) 73,540–549 547
 F. Rothhammer and T. D. Dillehay
Meltzer, D. J. (2006) Paleoamerican origins: Beyond clovis. J Field
Archaeology 31, 441–443.
Merriwether, D., Rothhammer, F. & Ferrell, R. (1995). Distribu-
tion of the four founding lineage haplotypes in native americans
suggests a single wave of migration for the new world. Amer J Phys
Anthropol 98, 411–430.
Merriwether, D. (2006) In: Handbook of North American Indians
Vol. 3: Environment, Origins and Populations (eds. D. Ubelaker,
W. C. Sturtevant & D. Stanford), pp. 817–830. Washington, DC:
Smithonian Institution Press.
Miotti, L. L., Salemme, M. & Flegenheimer, N. (2003). Where the
South Winds Blow: Ancient Evidence of Paleo-South Americans. Col-
lege Station, USA: Texas A&M University Press.
Morell, V. (1990) Confusion in earliest America. Science 248, 439–
441.
Neves, W. A. & Hubbe, M. (2005) Cranial morphology of early
Americans from Lagoa Santa, Brazil: implications for the settle-
ment of the New World. Proc Natl Acad Sci 102(51), 18309–18314.
Neves, W. A., Prous, A., Gonzalez-Jose, R., Kipnis, R. & Powell, J.
(2003) Early Holocene human skeletal remains from Santana do
Riacho, Brazil: implications for the settlement of the New World.
J Hum Evol 45, 19–42.
Neves, W. A. & Pucciarelli, H. M. (1990) The origin of the first
Americans: an analysis based on the cranial morphology of early
South American human remains. Am J Phys Anthropol 81, 274.
Neves, W. A. & Pucciarelli, H. M. (1991) Morphological affinities of
the first Americans: an exploratory analysis based on early South
American human remains. J Hum Evol 21, 261–273.
O’Rourke, D. H. & Suarez, B. K. (1985) Pattern and correlates of
genetic variation in South Amerindians. Ann Hum Biol 13, 13–32.
Piperno, D. R. & Pearsall, D. M. (1998) The Origins of Agriculture in
the Lowland Neotropics. New York: Academic Press.
Piperno, D. & Stothert, K. (2003) Phytolith evidence for early
holocene cucurbita domestication in southwest Ecuador. Science
299, 1054–1057.
Pietschmann, R. (1906) Geschichte des Inkareiches von Pe-
dro Sarmiento de Gamboa Abhandlungen der koeniglichen
Gesellschaft der Wissenschaften zu Goettingen. In: Philologisch-
historische Klasse. Berlin.
Powell, J. F. & Neves, W. A. (1999) Craniofacial morphology of the
first Americans: pattern and process in the peopling of the New
World. Am Phys Anthropol 42, 153–188.
Prieto, A. R. (1996) Late quaternary vegetational and climatic
changes in the Pampa grassland of Argentina. Quaternary Res 45,
73–88.
Prous, A. & Fogaça, E. (1999) Archaeology of the Pleistocene-
Holocene boundary in Brazil. Quat Int 53/54, 21–41.
Prugnolle, F., Manica, A. & Balloux, F. (2005) Geography pre-
dicts neutral genetic diversity of human populations. Curr Biol
15, R159-R160.
Pucciarelli, H. M. (2004) Migraciones y variación craniofacial hu-
mana en América. Complutum 15, 225–247.
Ramachandran, S., Dreshpande, O., Roesman, C. C., Rosenberg,
N. A., Feldman, M. W. & Cavalli-Sforza, L. L. (2005) Support
from the relationship of genetic and geographic distance in human
populations for a serial founder effect originating in Africa. Proc
Natl Acad Sci USA 102, 15942–15947.
Relethford, J. H. (2002) Apportionment of global human genetic di-
versity based on craniometrics and skin color. Am J Phys Anthropol
118, 393–398.
Rivet, P. (1909) Los Orı́genes del Hombre Americano. Mexico-Chile:
Fondo de Cultura Económica.
548 Annals of Human Genetics (2009) 73,540–549
Journal compilation 
Rivet, P. (1943) Recherches anthropologiques sur la Basse-
Californie. J Soc Amer Paris 6, 3–109.
Rothhammer, F. & Silva, C. (1992) Gene geography of Souh Amer-
ica:Testihg models of population displacement based on archeo-
logical evidence. Am J Phys Anthropol 89, 441–446.
Rothhammer, F., Silva, C., Callegari-Jacques, S. M., Llop, E. &
Salzano, F. M. (1997) Gradients of HLA diversity in South Amer-
ican Indians. Ann Hum Biol 24, 197–208.
Rull, V. (1996) Late pleistocene and holocene climates of Venezuela.
Quat Int 31, 85–94.
Salemme, M & Miotti, L. (2002) South America: long and wind-
ing roads for the first Americans at the Pleistocene/Holocene
transition. Editors Salemme, M, & Miotti, L. Quat Int 109–110,
1–179.
Sandweiss, D. H., McInnis, H., Burger, R., Cano, A., Ojeda, B.,
Paredes, R., Sandweiss, M. del C. & Lascock, M. (1998) Quebrada
Jaguay: Early South American maritime adaptations. Science 281,
1830–1832.
Stanford, D. & Bradley, B. (2002). Ocean trails and prairie paths?
Thoughts about Clovis Origins. In Jablonski, N., editor. The
First Americans: The Pleistocene colonization of the New World.
Memoirs of the California Academy of Sciences 27. 255–271.
Starikovskaya, E. B., Sukernik, R. I., Derbeneva, O. A., Volodko, N.
V., Ruiz-Pesini, E, Torroni, A., Brown, M. D., Lott, M. T., Hos-
seini, S. H., Huoponen, K. & Wallace, D. C. (2005) Mitochondrial
DNA diversity in indigenous populations of the southern extent
of Siberia, and the origins of Native American haplogroups. Ann
Hum Genet 69, 67–89.
Steele, D. G. & Powell, J. F. (1992) Peopling of the Americas: pale-
obiological evidence. Hum Biol 64, 303–336.
Stone, A. C. & Stoneking, M. (1998) mtDNA analysis of a pre-
historic Oneota population: implications for the peopling of the
New World. Am J Hum Genet 62, 1153–1170.
Stothert, K. (1998) An early holocene maritime adaptation in south-
west Ecuador: New perspectives on the Las Vegas evidence. Paper
presented at 63rd Ann. Meeting of the SAA, Seattle.
Szathmáry, E. J. E. (1993a) MtDNA and the peopling of the Amer-
icas. Am J Hum Genet 53, 793–799.
Szathmáry, E. J. E. (1993b) Genetics of aboriginal North Americans.
Evol Anthropol 1, 202–220.
Tarazona-Santos, E., Carvalho-Silva, D. R., Pettener, D., Luiselli,
D., De Stefano, G. F., Labarga, C. M., Rickards, O., Tyler Smith,
C., Pena, S. D. & Santos, F. R. (2001) Genetic differentiation
in South America is related to environmental and cultural di-
versity:evidence from the Y chromosome. Am J Hum Genet 68,
1485–1496.
Turner, C. G. (1987) Late Pleistocene and Holocene population
history of East Asia based on dental variation. Am J Phys Anthropol
73, 305–321.
Volodko, N. V., Starikovskaya, E. B., Mazunin, I. O., Eltsov, N. P.,
Naidenko, P. V., Wallace, D. C. & Sukernik, R. I. (2008) Mi-
tochondrial genome diversity in arctic Siberians, with particular
reference to the evolutionary history of Beringia and Pleistocenic
peopling of the Americas. Am J Hum Genet 82, 1084–1100.
Walter, H. V., Cathoud, A. & Mattos, A. (1937) The Confins man. A
contribution to the study of early man in South America. In: Early
Man: As depicted by leading authorities at the International Symposioum
of the Academy of Natural Sciences, Philadelphia, March 1937 (ed.
G. G. MacCurdy) London: Lippincott.
Wang, S., Lewis, C. M., Jakobsson, M., Ramachandran, S., Ray,
N., Bedoya, G., Rojas, W., Parra M. V., Molina, J. A., Gallo, C.,
Mazzotti, G., Poletti, G., Hill, K., Hurtado, A. M., Labuda, D.,

C 2009 The Authors
C 2009 Blackwell Publishing Ltd/University College London
 Late Pleistocene Colonization of South America

Klitz W., Barrantes, R., Bortolini, M. C., Salzano, F. M., Petzl-
Erler, M. L., Tsuneto, L. T., Llop, E., Rothhammer, F., Excoffier,
L., Feldman, M. W., Rosenberg, N. A. & Ruiz-Linares, A. (2007)
Genetic variation and population structure in native Americans.
PLoS Genet 3, e185.
Weng, C., Bush, M. B. & Silman, M. R. (2004) An analysis of
modern pollen rain on an elevational gradient in southern Peru.
J Trop Ecol 20, 113–124.
Wiesner, G. (1999) Early Ecuador people were maritime adapted.
Mammoth Trumpet 14, 4–11.
Zegura, S. L., Karafet, T. M., Zhivotovsky, L. A. & Hammer, M. F.
(2004) High-resolution SNPs and microsatellite haplotypes point
to a single, recent entry of Native American Y chromosomes into
the Americas. Mol Biol Evol 21, 164–175.
Received: 8 October 2008
Accepted: 24 June 2009


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