doi 10.4436/JASS.

91002 JASs Invited Reviews

e-pub ahead of print Journal of Anthropological Sciences
Vol. 92 (2014), pp. 79-97

Homo sapiens in the Americas. Overview of the earliest

human expansion in the New World
Aurelio Marangoni1, David Caramelli2 & Giorgio Manzi1

1)Dipartimento di Biologia Ambientale, Sapienza Università di Roma, Piazzale Aldo Moro 5, 00185
Roma, Italy
e-mail: giorgio.manzi@uniroma1.it

2) Dipartimento di Biologia Evoluzionistica, Università di Firenze, Via del Proconsolo 12, 50122 Firenze, Italy

Summary - Although it is widely recognised that America was the last continent to be populated by
our species, researchers’ views on various aspects of this process (e.g. the period in which it occurred, the
area from which the colonizing populations came, the number of dispersal waves and the routes taken by
these migrations) differ significantly. In this paper, we review both classical data and more recent findings
from various research fields – including geology, paleoecology, archaeology, skeletal biology, and genetics –
that may shed light on the dynamics of the colonization of the American continent, according to a critical
reappraisal of the various hypotheses and models that have been advanced over time to explain this process.

Keywords - Early modern humans, Migratory routes, Models of dispersal, Last Glacial Maximum,
America.

Introduction including the Jesuit Jose de Acosta (1589) and

the naturalist Georges-Louis Leclerc de Buffon
The dispersal of human populations in the (1749), who were among the first to speculate on
American continent may be viewed as a part of the origin of the ancestors of the Native American
the worldwide process of expansion of our species populations (Mazieres, 2011). Thus, in the 20th
which, as consistently demonstrated by fossil and century numerous hypotheses were advanced to
genetic data (e.g., Cann et al., 1987; Stringer & describe the first settlement of the Americas in
Andrews, 1988; Manzi et al., 1994; Harpending the light of archaeological, anthropological and
et al., 1998; Stringer, 2002; McDougall et al., (more recently) genetic data.
2008; Endicott et al., 2010; Manzi, 2012), first In this review, we consider both classical
appeared in Africa around 200 thousand years data and recent findings from various research
ago (200 ka). fields - such as geology, paleoecology, archaeol-
Homo sapiens started spreading out of Africa ogy, skeletal biology, and genetics – that have
between 120 and 60 ka, gradually reaching the been taken into account to explain the dynamics
landmasses found elsewhere on the planet at of the colonization of the American continent.
various times; the Americas were the last conti- We also briefly discuss the various hypotheses
nent to be inhabited in this expansion process and models that have been proposed over time
(Cavalli-Sforza & Feldman, 2003; Endicott et to describe this process, including: the Clovis-
al., 2003; Foster & Matsumura, 2005; Makaulay first/single origin hypothesis (Hrdlicka, 1937;
et al., 2005; Tangeraj et al., 2005; Mellars, 2006; Haynes, 2002; Fagan, 2004), the Blitzkrieg/
Rose et al., 2011). Ever since the 16th century, overkill model (Martin, 1973, 1984), the Three-
the colonization of the American continent wave migration/tripartite model (Greenberg et
has attracted the interest of many intellectuals, al., 1986), the Solutrean hypothesis (Bradley &

the JASs is published by the Istituto Italiano di Antropologia www.isita-org.com

 80 The colonization of the American continent

Stanford, 2004), the Two main biological com- a wide plain free of ice, containing a rich variety
ponents/stock hypothesis (Neves & Pucciarelli, of plant species, herds of herbivorous mammals
1991; Pucciarelli et al., 2003), the Beringian (mammoths, musk oxen, horses, camels, bisons)
incubation/source population model (Szathmáry, as well as their predators (lions, short-faced bears,
1981, 1996; Foster, 1996; Bonatto & Salzano, sabertoothed cats) (Guthrie, 2001).
1997; Bodner, 2012), the Three-stage coloni- Moreover, stone artefacts used on the skel-
zation model (Kitchen et al., 2008, Mulligan etal remains of extinct mammals have recently
et al., 2008), the Single migration hypothesis been described at Yana Rhinoceros Horn (north-
(Fagundes et al., 2008), the Dual migration western Siberia), demonstrating that Beringia
hypothesis (Torroni, 1992; Schurr & Sherry, was inhabited by human groups from at least
2004; Perego et al., 2009), and the Recurrent 32 ka (Pitulko et al., 2004). In recent years,
gene flow model (Ray et al., 2009; De Azevedo other archaeological sites have been described
et al., 2011 ). Nevertheless, we believe that such in both Alaska and eastern Siberia; these sites
a puzzling anthropological issue may be better include: i) Swan Point (central Alaska), where
understood by combining a multidisciplinary artefacts consisting of both microblades and bur-
approach with a large scale genetic analysis, as a ins dated to 14 ka were found; ii) Nenana (cen-
recent study (Reich et al., 2012) suggests. tral Alaska) and Ushki (south-western Siberia),
which yielded small blades and flakes dated to
between 13.8 and 13 ka; iii) Sluiceway-Tuluaq
At the end of the Pleistocene (north-western Alaska), where lanceolate bifaces
dated to between 13.4 and 13 ka were brought
The paths that may have been followed by to light; and (iv) Nohagabara I (western Alaska),
the human populations that first colonized where bifaces and microblades cores dated to
the Americas were influenced by the climatic between 13.8 and 12.7 ka were found (reviewed
and environmental effects of the last glaciation in Goebel et al., 2008; Dillehay, 2009).
(Marine Isotopic Stages 4-2), which occurred However, human populations were presum-
during the Late Pleistocene (Guthrie, 2001; ably unable to reach southern Alaska, during the-
Mandryk et al., 2001, Peltier, 2002; Meltzer, extensive marine regression that characterized the
2009; Dixon, 2011). LGM, because the Laurentide and Cordilleran gla-
In particular, during the Last Glacial ciers formed a continuous ice mass that extended
Maximum (LGM) around 21 ka (Clark & Mix, from the Pacific to the Atlantic coasts (Meltzer,
2002), most of present-day Canada and the 2009). Paleoclimate studies have shown that the
northern United States were covered by two first accessible route for human populations was
main glaciers: the Laurentide glacier (which probably along the Pacific coast, which became free
covered approximately 13.4 million km2 and of ice around 15 ka, while an inland passage (“ice-
extended from the Atlantic coast to the foothills free corridor”) opened when the glaciers began
of the Rocky Mountains) and the Cordilleran to separate at the foot of the Rocky Mountains,
glacier (which covered about 2.4 million km2 and though this did not probably occur until 13.5 ka
extended from the Pacific coast to the western (Mandryk et al., 2001; Dixon, 2011).
edge of the Rocky Mountains) (Meltzer, 2009).
During the coldest phases of the Late Pleistocene
the sea level was as much as 120 meters lower The archaeological evidence
than it is today (Peltier, 2002), and Asia and
America were connected by a land bridge known The first archaeological evidence of prehis-
as Beringia, corresponding to what is now the toric human settlements in the Americas was dis-
Bering Strait (Fig. 1). Fossil records indicate covered in 1927 at Folsom in New Mexico, where
that, around the time of the LGM, Beringia was some spearheads associated with the remains of a
 J. Hawkset al.
A. Marangoni 81

Fig. 1 - Map showing the main physical characteristics of the American continent during the Last
Glacial Maximum (LGM), with a selection of the prehistoric sites mentioned in the text. The ice-free
corridor opened approximately after 13.5 ka. Adapted from Meltzer (2009).

Late Pleistocene bison were found, and in 1933 proved to have been the most widespread cul-
at Clovis, which is also in New Mexico, where tural tradition on the continent, though more
other less sophisticated, and therefore prob- recent dates obtained for several Clovis sites
ably earlier, spearheads were discovered near the have shown that it only spanned approximately
remains of a mammoth (Howard, 1933). In the 400 years, from 13.2 to 12.8 ka (Waters Jr. &
following years lithic artefacts, made with what Stafford, 2007), after which it was replaced by
was to become known as Clovis technology, were other cultural traditions.
found in many archaeological sites in North and The typical elements of the Clovis culture
Central America. The Clovis technology thus are fluted projectile points, made with siliceous

www.isita-org.com
 82 The colonization of the American continent

rocks such as chert, jasper, chalcedony and obsid- mammals, in both North and South America
ian, modelled by removing large blades from a (Greyson & Meltzer, 2003), some researchers
nucleus and refinished so as to obtain a laminar believe that the extinction of these species was
shape that is tapered on either side (Goebel et al., the result of hunting on a massive scale practised
2008). by the Clovis people, in what has been called the
Following the discovery of Clovis artefacts, Blitzkrieg/overkill model (Martin, 1973, 1984).
it was assumed for many decades that the first However, this assumption has been repeat-
people to reach the American continent were edly questioned in recent years because there
hunter-gatherer groups using this technology, is no archaeological evidence of the killing of
who had come from Asia at the end of the LGM animals other than mammoths and mastodons
and had populated the entire continent, from the during the Clovis age, and because both the Late
Hudson Bay to Tierra del Fuego, over a period Pleistocene and the Pleistocene/Holocene transi-
of a thousand years: a scenario referred to as the tion are characterized by numerous extinctions,
Clovis-first/single origin hypothesis (Hrdlicka, even in regions in which human populations
1937; Haynes, 2002; Fagan, 2004). are known not to have been present (Greyson &
However, while artefacts produced with the Meltzer, 2003).
Clovis technology have been described in many Whether or not it was due to anthropogenic
North American archaeological sites, the archaeo- causes, the extinction of these species is very likely
logical record for South America is very differ- to have had an important impact on the history of
ent, with no single material culture dominating the American continent. Indeed, according to the
this sub-continent. South American sites that are hypothesis that human infectious diseases evolved
contemporary with the Clovis phase are charac- as the result of the interaction between humans
terized by a marked degree of regional cultural and domestic animals (Diamond, 2000), Native
diversification, with at least six sites being iden- Americans populations were deprived of the
tified (Cerro Tres Tetas, Cueva Casa del Minero opportunity to develop any immunity over time
and Piedra Museo in Argentina, and Fell’s Cave, owing to the rapid disappearance of many poten-
Quebrada Santa Julia and Quebrada Jaguay in tially domestic species. When Native Americans
Chile) in which a wide range of flake tools, bifa- subsequently came into contact with Europeans
cial points and, in some cases, distinctive Fishtail in the late 15th century, they suffered a high mor-
points have been described (reviewed in Goebel tality caused by infectious diseases such as small-
et al., 2008). Some authors have ascribed this cul- pox, tuberculosis, measles, rubella and syphilis,
tural diversity to geographical barriers that ham- as archaeological and historical records indicate.
pered human movements (such as the Andean Moreover, a recent genetic study (O’Fallon &
highlands and Amazon river) as well as to climatic Fehren-Schmitz, 2011) indicates that Native
and ecological changes between areas at different American populations were subject to a signifi-
latitudes (Rothhammer & Dillehay, 2009). cant contraction in population size around 500
years ago, when the number of females is known
to have decreased by around 50%.
Late Pleistocene extinctions

Clovis spearheads have frequently been asso- In search of greater antiquity

ciated with skeletal remains of proboscideans,
there being at least 14 sites dating from the The identification of pre-Clovis archaeo-
Clovis period in which mammoths and masto- logical sites proved to be an arduous task owing
dons were killed and butchered (Meltzer, 2009). to the difficulty encountered in many cases in
Since the American Late Pleistocene is charac- understanding whether chipped stones and
terized by the extinction of 35 genera of large crushed mammal bones were the result of human
 J. Hawkset al.
A. Marangoni 83

activity or were merely due to natural processes do Riacho and Lapa do Boquete (Brasil), dat-
(Lyman et al., 1998). In other cases, establish- ing from around 14 ka, and at Taima-Taima
ing the age of a site proved to be difficult owing (Venezela), dating from 13.2 ka (reviewed in
to the lack of reliable radiometric dates (Meltzer, Dillehay, 2009).
2009). In view of these difficulties, many archae- In brief, as we shall see below, the body of
ological sites that had until the mid-1990s been evidence yielded by different fields of research
considered to point to an early colonization of allows us to state with some confidence that the
the Americas (approx. 50-30 ka), such as those hunter-gatherers who developed the Clovis tech-
at Calico (California), Pedra Furada (Brazil), nology were not the first inhabitants of America
Pendejo Cave (New Mexico), Pikimachay Cave (Waters Jr. & Stafford, 2007).
(Peru), Tlapacoya (Peru) and Tule Springs
(Nevada) (reviewed in Goebel et al., 2008), are
no longer considered a robust evidence and are Looking for a multidisciplinary
generally disregarded. approach
The first archaeological site accepted by
the vast majority of researchers as pre-Clovis is The first attempt to use a multidiscipli-
Monte Verde in southern Chile, dated to 14.6 nary approach to describe the earliest arrival of
ka, where the exceptional state of preservation humans in the Americas dates back to the late
of organic materials, interpreted as the remains 1980s and is known as the Three-wave migra-
of a human encampment, provided a precious tion/tripartite model (Greenberg et al., 1986).
insight into the daily lives of the inhabitants of This model is mainly based on studies by the
the Americas at the very end of the Pleistocene linguist Joseph Greenberg who, upon comparing
(Dillehay, 1997; Dillehay et al., 2008). the languages spoken by Native American groups,
Many other pre-Clovis sites have been grouped them into three families: Amerindian,
described in recent years in North America. Na-dene (or Athabascan) and Eskimo-Aleut.
Those generally accepted by the majority of According to Greenberg, the populations of each
authors include: i) the Debra L. Friedkin Site of these language families were descended from
(Texas), where more than 15,000 artefacts, con- three groups that had reached the Americas at
sisting mainly of small blades, and a block of different times: the Amerindians around 11 ka,
hematite, probably used to obtain red pigment, the Na-Denes 9 ka and the Eskimo-Aleuts 4 ka
dating from between 15.5 and 13.2 ka, were dis- (Greenberg et al., 1986) (Fig. 2a).
covered (Waters et al., 2011); ii) Meadowcraft On the basis of the frequency of 28 dental
Rockshelter (Pennsylvania), which contained traits spread among Native American populations,
stone tools dating from between 15.2 and the physical anthropologist Christy Turner II also
13.4 ka, but that may be even older (Adovasio came to the conclusion that the early settlers of the
& Pedler, 2004); iii) Schaefer and Hebior Americas could be subdivided into three groups,
(Wisconsin), with stone artefacts and skeletal i.e. the Eskimo-Aleuts, Greater Northwest Coast
remains of mammoth with cutmarks, dating Indians and Macro-Indians. According to Christy
from between 14.5 and 14.2 ka (Joyce, 2006); Turner II, the Macro-Indians (who corresponded
iv) Page-Ladson (Florida), dating to 14.4 ka, to the Amerindians of Greenberg) had reached
where artefacts associated with bones of extinct America around 14 ka; the order of the other
Pleistocene animals, including a mastodon tusk two groups instead differed from Greenberg’s
with cutmarks where it was joined to the skull, order insofar as Christy Turner II considered the
were found (Webb, 2005). Eskimo-Aleuts to have arrived about 11 ka, and
Other archaeological evidence, besides that the Greater Northwest Coast Indians (who only
found at Monte Verde, pre-dating the Clovis partially corresponded to the Na-denes) to have
period exists in South America: at Santana reached America last (Greenberg et al., 1986).

www.isita-org.com
 84 The colonization of the American continent

Fig. 2 – a) According to the Three-wave migration/tripartite model, which combined linguistic, den-
tal and genetic evidence, Native Americans descended from three groups of migrants belonging to
distinct linguistic families: Amerindians, Na-denes and Eskimo-Aleutians. b) The Single migration
hypothesis states that America was settled through one migration route along the Pacific coast,
whereas two independent migration routes were followed according to the Dual migration hypoth-
esis. In addition, some genetic models suggest that colonization from Asia was interrupted by a long-
term settlement in Beringia, as originally pointed out by E. Szathmáry (1981).
 J. Hawkset al.
A. Marangoni 85

When the geneticist Stephen Zegura joined America, they speculated that people had crossed
Greenberg and Turner II, the genetic analyses the Atlantic from Spain during the LGM, when
available at the time were based exclusively on North America and Europe were connected by
classical genetic markers (blood groups, serum a bridge of land and continental glaciers. They
proteins, enzyme polymorphisms) (Cavalli- claimed that their hypothesis was supported by
Sforza et al., 1994, among others), whose reso- the fact that the only archaeological sites con-
lution was not sufficient to separate Native sidered at the time to be of pre-Clovis age – i.e.
Americans into different groups. As the only Meadowcroft Rockshelter (Pennsylvania), Cactus
marker that appeared to confirm the three-wave Hill (Virginia) and Page Ladson (Florida) – were
migration model was the distribution of the allo- all located on the eastern side of North America
types of the immunoglobulin G (Williams et al., (Bradley & Stanford, 2004).
1985), Zegura concluded that the genetic data Most researchers now consider the similarity
available could not be used to support the results between the Solutrean and Clovis lithic technol-
yielded by the linguistic and dental analyses ogies to be due to cultural convergence (Straus et
(Greenberg et al., 1986). al., 2005); moreover, genetic studies carried out
on Native American populations have revealed
their close affinity with Asian populations, which
Colonization from Europe? appears to disprove the Solutrean hypothesis
(Schurr et al., 1990; Torroni et al., 1993; Karafet
When the Clovis artefacts first appeared, et al., 1999).
numerous archaeologists were struck by their
resemblance to those of the Solutrean culture
of the European upper Paleolithic, dated to Skeletons of first Americans
between 23.5 and 20 ka (Straus et al., 2005).
For example, the archaeologist Arthur Jelinek Human skeletal remains dating from the
(1971) described the artefacts from the Franco- Late Pleistocene and the Early Holocene – thus
Cantabrian area as being highly similar to the closely related to the earliest inhabitants of the
Clovis artefacts, although a historical relation- Americas – have so far only been found in sites
ship between these two technologies was, in his located in a limited number of regions on the
view, unlikely because they are separated tempo- continent, most of which lie in South America
rally by more than 6,000 years and geographi- (Jantz & Owsley, 2001). These sites are: i)
cally by the Atlantic Ocean. Kennewick (Washington), dated to 9 ka (Taylor
In 2004, however, the experimental archaeol- et al., 1999), and Warm Mineral Spring (Florida),
ogists Bruce Bradley and Dennis Stanford, hav- dated to 10 ka (Powell et al., 1999), in the USA;
ing probably been influenced by the discovery in ii) the Mexico basin, dated to 10 ka (Gonzales-
North America of some individuals with the X Jose et al., 2005), and Baja California, dated to
haplogroup of mitochondrial DNA (mtDNA), 6 ka (Gonzales-Jose et al., 2003), in Mexico; iii)
which also occurs in European populations Sabana de Bogotà, dated to between 10 and 6
(Brown et al., 1998), argued that such a hypoth- ka (Neves & Pucciarelli, 1991), in Colombia;
esis deserved to be reconsidered (Meltzer, 2009). iv) Toca de Oncas, dated to 8 ka (Hubbe et al.,
Bradley and Stanford founded their Solutrean 2004), Lagoa Santa, dated to between 11 and
hypothesis on the view that, despite being highly 7.5 ka (Neves & Hubbe, 2005), and Capelinha,
complex, the Clovis lithic technology cannot be dated to 8.5 ka (Neves et al., 2005), in Brazil; v)
traced to other technologies in either Siberia or Arroyo Seco 2, dated to between 9.8 and 8.3 ka
Alaska but is, in their view, closely related to the (Scabuzzo & Politis, 2007), in Argentina; and vi)
Solutrean technology. In order to explain how Palli Aike, dated to between 8 and 7 ka (Neves et
some European populations might have reached al., 1999), in Chile.

www.isita-org.com
 86 The colonization of the American continent

When researchers analysed craniometric view the groups usually labelled Paleoamericans,
variation in both ancient and recent Native Amerindians and Mongoloids as variants along
Americans, using traditional methods at first and a morphological continuum. In another recent
geometric morphometrics more recently, they study, based on both cranial morphology and
found morphological differences that led to the molecular data, Perez and colleagues (2009)
overall sample being divided into two groups. analysed skeletal samples from central-eastern
Assuming that cranial morphology reflects Argentina (Arroyo Seco 2), where a diachronic
the history of human populations (Neves & sequence ranging from Early to Late Holocene
Pucciarelli, 1991), this difference has been inter- occurs. Although the morphology of the dia-
preted by some as evidence of at least two sepa- chronic samples – i.e. those from i) the Middle
rate migratory events in the American continent and Late Holocene sample and ii) from the other,
over time. more recent Late Holocene sample – is clearly
The crania of the earliest Americans (earlier different, they found that both groups belong
than around 10 ka) have been referred to as the to the same mtDNA haplogrups. It has conse-
“Paleoamericans”. These tend to be similar to quently been suggested that the morphological
those of extant Australians, Melanesians and Sub- variations between ancient and recent Native
Saharan Africans, like the Late Pleistocene speci- Americans is more likely the result of random
men from Zhoukoudian Upper Cave (skull 101; (genetic drift) or non-random (natural selection
Neves & Pucciarelli, 1998). Indeed, they exhibit a or plasticity) micro-evolutionary factors than of
dolichocephalic morphology, prognathic and low distinct waves of immigrants.
faces, with relatively low, broad orbits and noses.
By contrast, late prehistoric and recent Native
Americans, also known as “Amerindians”, tend to Genetic evidence
exhibit a cranial morphology similar to later and
modern East Asian populations (“Mongoloids”), Autosomal genes, Y chromosome, mitochondrial
with a brachymorphic cranial vault, orthognathic DNA
high, broad faces, and relatively high and narrow Many Native American populations have
orbits and noses (Lahr et al., 1995; Van Vark et been genetically characterized since the second
al., 2003; Neves & Hubbe, 2005). half of the 20th century. The characterization of
According to this two-wave model, referred these populations was initially based on classic
to as the Two main biological components/ genetic markers [i.e. on the expression of autoso-
stock hypothesis (Neves & Pucciarelli, 1991; mal markers at the blood level (red cell, protein
Pucciarelli et al., 2003), the Paleoamericans and serum systems) (Cavalli-Sforza et al., 1994;
derived from South-East Asian populations of Crawford, 1998; O’Rourke, 2006)], whereas in
the Late Pleistocene, which spread throughout the late 1990s both maternal (mitochondrial
America around 16 ka, whereas the Amerindians DNA) and paternal (non-recombining por-
derived from the East Asian populations of tion of Y chromosome, or NRY) markers were
the Early Holocene, which spread through the included in the analyses.
American continent around 10 ka, replacing, As regards the autosomal genes, Wang and
or assimilating, the Paleoamericans (Lahr et al., colleagues (2007) recently studied 24 Native
1995; Van Vark et al., 2003; Neves & Hubbe, American populations: not only did they show
2005; Hubbe et al., 2010; Hubbe et al., 2011). that the average level of heterozygosity of Native
Many scholars have criticized this scenario: American autosomal genes is 6.5% lower than
for example, Gonzales-Jose and colleagues the average of our species, but they also found a
(2008), after analysing a group of 576 Late decline in genetic variability that was directly cor-
Pleistocene/Early Holocene and modern crania, related with the population’s distance from the
suggested that it would be more appropriate to Bering Strait. When Schroeder and colleagues
 J. Hawkset al.
A. Marangoni 87

(2009) studied a D9S1120 allele with a high Ancient DNA

frequency in Native American populations, they The analysis of samples of genetic material
argued that it was more likely to have resulted taken directly from ancient biological remains
from common descent within a small founder (ancient DNA analysis), despite certain limita-
population than from natural selection. tions (i.e. limited availability of samples, pres-
Studies on the NRY, including the analysis ervation state of organic material, possible con-
of both the Single Nucleotide Polymorphisms tamination), may be considered the only means
(SNPs) and Short Tandem Repeats (STRs), of directly investigating the genetic diversity of
have shown that the two most common hap- extinct populations.
logroups in Native American populations are The earliest hard evidence (i.e. not merely
Q and C, while the most common lineages are archaeological) of the presence of humans on the
Q1a*-MEH2, Q1a3*-M346, Q1a3a1-M3 and American continent came to light in 2008 at the
C3b-P39 (Karafet et al., 2006; Karafet et al., Paisley 5 Mile Point Caves in Oregon, where 14
2008; Zegura et al., 2004; Blanco-Vetea et al., human coprolites, dated to 14.27-14 ka, were
2010; Jota et al., 2011). Mulligan and colleagues discovered among the remains of an encamp-
(2004) and Zegura and colleagues (2004) have ment. The DNA obtained from 6 of these copr-
estimated that the coalescence for both these olites yielded Native American SNPs diagnostic
haplogroups occurred between 12 and 15 ka. for the mtDNA founding haplotypes A2 and B2
Moreover, a high-resolution analysis of the hap- (Gilbert et al., 2008a).
logroup Q, partially reshaped the phylogeny of A biological sample (teeth) from the On-Your-
this branch and showed that Native Americans Knees Cave site on the Prince of Wales Island in
and Southern Altaian populations share a com- Alaska, dated to 10.3 ka, proved to belong to an
mon ancestor (Dulik et al., 2012), thereby pro- individual characterized by the mtDNA linege
viding important information concerning the D4h3a (Kemp et al., 2007): a rare lineage that has
possible origin of the populations that colonized been found in some extant individuals living on the
the American continent. Pacific coast of South America (Perego et al., 2007).
Early research on mtDNA, using both Genetic material was also recently obtained
Restriction Fragment Length Polymorphisms from a frozen hair belonging to a Paleo-Eskimo
(RFLPs) and sequencing of the hypervariable individual excavated from an early settlement in
segment I (HVR-I), indicated that all Native Greenland (Saqquaq) and dated to 4.5-3.4 ka.
Americans belonged to five haplogroups (A, B, The sequencing of 79% of the nuclear genome
C, D and X) (Torroni et al., 1993; Forster et al., showed this individual possessed thick, dark hair,
1996; Brown et al., 1998). A greater degree of brown eyes and probably shovel-shaped front
molecular resolution, achieved in recent years teeth, a characteristic that is common among
by sequencing entire mtDNA (Bandelt et al., both Asian and Native American populations
2003; Achilli et al., 2008), led to the identifica- (Rasmussen et al., 2010). The Seqquaq Paleo-
tion of 15 sub-haplogroup lineages considered Eskimo was also found to belong to the mtDNA
to be founders (A2*, A2a, A2b, B2, C1b, C1c, lineage D2a1, a lineage that had not previously
C1d*, C1d1, C4c, D1, D2a, D3, D4h3a, X2a been observed among modern Native American
and X2g) (Perego et al., 2010). Although there and Neo-Eskimo populations. This indicates that
are some discrepancies in the coalescence age the first humans to colonize the far north of the
estimates of the mtDNA haplogroups between American continent were not either the Native
studies, they commonly range from 15 to 20 ka Americans or the ancestors of the Neo-Eskimo
(O’Rourke & Raff, 2010). This date is a rough expansion (Thule culture), which instead started
approximation of the drastic genetic bottleneck in Alaska around 1 ka (Gilbert et al., 2008b).
that forms the basis of one hypothesis regarding Moreover, the genetic characterization of
the earliest colonization of the Americas. samples belonging to two individuals from the

www.isita-org.com
 88 The colonization of the American continent

China Lake site in British Columbia, dated to and Asian individuals. They hypothesised, on
4.95 ka, showed that these individuals belonged the basis of the phylogenetic structure of the
to the mtDNA haplogroup M: an Asian hap- mtDNA haplogroups of Native Americans, that
logroup no longer present in Native American this ancestral population, upon being prevented
populations (Malhi et al., 2007). This is the first, from heading further South by the last glaciation,
and so far only, reported case of genetic discon- was forced to settle in Beringia for such a long
tinuity between ancient and modern Americans. time (Beringian standstill) that the New World
As very few biological remains with good founder lineages had sufficient time to differenti-
nucleic acid preservation have been found in ate from their Asian sister clades (Fig. 2b).
archaeological sites, the vast majority of ancient Kitchen and colleagues (2008) and Mulligan
DNA studies on ancient American populations and colleagues (2008) instead used the Bayesian
were conducted on samples younger than 5 skyline plot analysis (Drummond et al., 2005)
ka. More recent samples show that haplogroup to develop a model used to explain the dynam-
frequencies for the populations of the entire ics of the colonization of the Americas, which
American continent have not changed substan- they called Three-stage colonization model.
tially over the last 4 ka, with similar haplogroups According to this model, the first phase in this
being obtained for both the ancient and modern process was the genetic divergence between peo-
populations, though some changes in regional ples of central-eastern Asia and the ancestors of
haplogroup patterns do emerge (reviewed in Raff Native Americans, which took place more than
et al., 2011). 30 ka; this phase was followed by a period of iso-
lation, spanning 7-15 ka, during which a genetic
drift process led to variants that later became
Models for the earliest Americans characteristic of Native American populations;
the third and final phase was the colonization of
In the last couple of decades, genetic analyses, the whole continent, which started about 16 ka
often combined with paleoecological and archaeo- and coincided with the end of the last Ice Age.
logical data, have allowed researchers to develop Some scholars have used the analysis of uni-
population models designed to interpret the ori- parental genetic markers to trace the possible
gin, dynamics and migratory routes followed by migratory routes followed by the first American
the first American settlers. settlers. For instance, according to Fagundes and
The first attempt to use uniparental colleagues (2008), the colonization of Beringia
genetic data to develop a model for the earli- during the LGM was characterized by a marked
est Americans was made by Bonatto & Salzano drop in the number of people migrating from
(1997); they analysed the mtDNA region of 544 Asia. By contrast, the end of the last Ice Age wit-
extant Native American individuals by testing nessed an increase in the number of people arriv-
the model developed by E. Szathmáry, who had ing from Asia, which coincided with the coloni-
used classical genetic markers (Szathmáry, 1981, zation of the continent and is hypothesized to
1984). It is known as the Beringian incubation have followed a trajectory along the Pacific coast
model or Beringian source population (Foster, between 18 and 15 ka, according to the so-called
1996; Bonatto & Salzano, 1997; Bodner 2012). Single migration hypothesis (Fig. 2b).
According to this model, Beringia played a cen- Other authors, such as Schurr and Sherry
tral role as the place in which the ancestral popu- (2004), instead support the scenario proposed
lation of the Americas differentiated genetically by Torroni and colleagues on the basis of genetic
before colonizing the continent. This model was data (1992), according to which the popula-
subsequently further elaborated by Tamm and tions known at that time as the Amerindians and
colleagues (2007), who analysed the complete Na-denes derived from independent waves that
mtDNA sequence of 623 Native Americans colonized the Americas via two migratory routes.
 J. Hawkset al.
A. Marangoni 89

The first route was along the Pacific coast and was the initial settlement of the American continent
followed around 14.7 ka by populations charac- after the LGM was followed by an extended gene
terized by high frequencies of the A, B, C and D flow between Asian and American populations
mtDNA haplogroups as well as by some peculiar inhabiting the Arctic landscapes. They called
NRY haplogroups. The other route was through this scenario the Recurrent gene flow model.
the ice-free corridor between the Laurentide and De Azevedo and colleagues (2011) support this
Cordilleran ice sheets (cf. Figs 1, 2b), which was model and believe that it could explain not only
taken by populations characterized by high fre- the extant genetic diversity, but also the variation
quencies of the X mtDNA haplogroup as well as in craniofacial shape observed between Native
other peculiar NRY haplogroups around 12.5 ka. American skeletons dating from different periods
Perego and colleagues (2009), who stud- in the past.
ied the geographical distribution of two rare
mtDNA lineages (D4h3 and X2a), came to the
same conclusion. They argue that the Americas Final remarks
were first colonised about 15-16 ka by two
human groups that followed these different As we have seen, many hypotheses have
routes: the Pacific coast and the ice-free corridor been advanced over time to explain the human
in the North. This hypothesis is supported by colonization of the American continent. Very
the recent identification of 14 living individuals recently, in the most comprehensive survey of
belonging to the extremely rare Native American genetic diversity in Native American popula-
mtDNA lineage C4c, who display the same age tions conducted so far, a large group of schol-
and geographical distribution as the mtDNA ars analysed data from 52 Native American
lineage X2a and appear to belong to the human and Eurasian groups, genotyped using single
group that entered North America from Beringia nucleotide polymorphisms. Their data indicate
through the ice-free corridor (Kashani et al., that the vast majority of Native American pop-
2012), as well as by the identification of 43 sub- ulations descend from a homogeneous group,
jects belonging to the rare mtDNA clades D1g which they called First American, that reached
and D1j (Bodner et al., 2012) and 46 belonging the Americas from Asia presumably by crossing
to the clades B2i2 (former B2l) and C1b13 (de the Bering Strait more than 15 ka; however, the
Saint Pierre et al., 2012a; de Saint Pierre et al., populations of the Arctic regions (speakers of
2012b). These haplogroups characterize popula- the Eskimo-Aleut languages) inherited almost
tions from the Southern Cone of South America half their ancestry from a second stream of Asian
(Chile and Argentina), whose geographical dis- genes, while the Chipewyan group from Canada
tribution and inferred origin are consistent with (speakers of the Na-Dene language) inherited
the earliest archaeological sites in South America, roughly one-tenth of their ancestry from a third
who may have taken the migratory route that ran stream (Reich et al., 2012) (Fig. 3).
along the Pacific coast. This scenario interestingly resembles the
By contrast, Ray and colleagues (2009), who Three-Wave migration/tripartite model proposed
used Approximate Bayesian computation (ABC) in the late 1980s which combined linguistic, den-
methods (Beaumont et al., 2002) to analyse 401 tal morphology and low resolution genetic data
autosomal microsatellite loci belonging to 29 (Greenberg et al., 1986). It however partially dif-
Asiatic and Native American populations, claim fers from the 1980s model insofar as Greenberg
that neither a single nor a dual wave of migra- and colleagues did not explicitly predict the pos-
tion from Asia can explain the observed level sible admixture between the First Americans and
of genetic diversity of the Amerindian popula- the subsequen t streams of Asian migrants.
tions. They believe, instead, that the data avail- Whether or not this scenario is confirmed
able may be better explained by assuming that by future studies, it seems reasonable to presume

www.isita-org.com
 90 The colonization of the American continent

Fig. 3 - Simplified reproduction of a neighbour-joining tree relating Native American to selected non-
American populations, based on Fst distances (Reich et al., 2012) and consistent with the Three-
wave migration/tripartite model introduced by Greenberg, Turner II & Zegura (1987). Modified
from Reich and co-workers (2012).

that further insights on the puzzling anthropo- archaeology, palaeoecology and linguistic data)
logical issue of the colonization of the New World combined with large scale genetic analyses (using
will most likely come from a multidisciplinary both recent and ancient DNA data) made pos-
approach (based on skeletal/dental morphology, sible by recent technological advances.
 J. Hawkset al.
A. Marangoni 91

Acknowledgements Americans: novel insights from South America’s

Southern Cone. Genome Res., 22: 811-820.
We are grateful to Guido Barbujani (University Bonatto S.L. & Salzano F.M. 1997. A single
of Ferrara) for reading a previous version of the and early migration for the peopling of the
paper and providing useful comments. We also wish Americas supported by mitochondrial DNA
to thank the editor and three anonymous review- sequence data. Proc. Natl. Acad. Sci. U.S.A., 94:
ers of the JASs, whose suggestions greatly improved 1866-1871.
the original manuscript. Fabio Di Vincenzo, Mary Bradley B. & Stanford D. 2004. The North
Anne Tafuri and Maria Luana Belli kindly pro- Atlantic ice-edge corridor: a possible Palaeolithic
vided constant help and support. Lewis Baker route to the New World. World Archaeol., 36:
revised the English manuscript. 459-478.
Brown M.D., Hosseini S.H., Torroni A., Bandelt
H.J., Allen J.C., Schurr T.G., Scozzari R.,
References Cruciani F., Douglas C. & Wallace D.C. 1998.
MtDNA haplogroup X: an ancient link between
Achilli A., Perego U.A., Bravi C.M., Coble M.D., Europe/Western Asia and North America? Am.
Kong Q.P., Woodward S.R., Salas A., Torroni J. Hum. Genet., 63: 852-861.
A. & Bandelt H.J. 2008. The phylogeny of the Cann R.L., Stoneking M. & Wilson A.C. 1987.
four pan-American mtDNA haplogroups: im- Mitochondrial DNA and human evolution.
plications for evolutionary and disease studies. Nature, 325: 31-36.
Plos One, 3: e1764. Cavalli-Sforza L.L., Menozzi P. & Piazza A.
Adovasio J.M. & Pedler R.D. 2004. Pre-Clovis 1994. The history and geography of human genes.
sites and their implications for human occupa- Princeton University Press, Princeton.
tion before the Last Glacial Maximum. In D.B. Cavalli-Sforza L.L. & Feldman M.W. 2003. The
Madsen (ed): Entering America: Northest Asia application of molecular genetic approaches to
and Beringia before the Last Glacial Maximum, the study of human evolution. Nature Genet.,
pp. 139-158. University of Utah Press, Salt 33: 266-275.
Lake City. Crawford M.H. 1998. The origins of Native
Bandelt H.J., Herrnstadt C., Yao Y.G., Kong Americans. Evidence from anthropological genet-
Q.P., Kivisild T., Rengo C. & Scozzari R. 2003. ics. Cambridge University Press, Cambridge
Identification of Native American founder (U.K.).
mtDNAs through the analysis of complete De Azevedo S., Nocera A., Paschetta C., Castillo
mtDNA sequences: some caveats. Ann. Hum. L., Gonzalez M. & Gonzalez-Jose R. 2011.
Genet., 67: 512-524. Evaluating microevolutionary models for the
Beaumont M.A., Zhang W.Y., Balding D.J. 2002. early settlement of the New World: the impor-
Approximate Bayesian computation in popula- tance of recurrent gene flow with Asia. Am. J.
tion genetics. Genetics, 62: 2025-2035. Phys. Anthropol., 146: 116-129.
Blanco-Vetea A., Jaime J.C., Brion M. & De Saint Pierre M., Bravi C. M., Motti J.M.B.,
Carracedo A. 2010. Y-chromosome lineages in Faku N., Tanaka M., Llop E., Bonatto S.L. &
Native South American population. Forensic Moraga M. (2012a) An alternative model for
Sci. Int. Genet., 4: 187-193. the early peopling of southern South America
Bodner M., Perego U.A., Huber G., Fendt L., revealed by analyses of three mitochondrial
Rock A.W., Zimmermann B., Olivieri A., DNA haplogroups. Plos One, 7: e43486.
Gomez-Carballa A., Lancioni H., Angerhofer De Saint Pierre M., Gandini F., Perego U.A.,
N., Bobillo M.C., Corach D., Woodward S.R., Bodner M., Gomez-Carballa A., Corach D.,
Salas A., Achilli A., Torroni A., Bandelt A.J. & Angerhofer N., Woodward S.R., Semino O.,
Parson W. 2012. Rapid coastal spread of First Salas A., Parson W., Moraga M., Achilli A.,

www.isita-org.com
 92 The colonization of the American continent

Torroni A. & Olivieri A. 2012b. Arrival of Fagundes N.J.R., Kanitz R., Eckert R., Valls
paleo-indians to the southern cone of South A.C.S., Bogo M.R., Salzano F.M., Smith
America: new clues from mitogenomes. Plos D.G., Silva Jr W.S., Zago M.A., Ribeiro-do-
One, 12: e51311. Santos A.K., Santos S.E.B., Petzl-Erler M.L.
Diamond J. 2000. Guns, germs and steel. The fate & Bonatto S.L. 2008. Mitochondrial popula-
of human societies. Norton & Company, New tion supports a single pre-Clovis origin with a
York. coastal route for the peopling of the Americas.
Dillehay T.D. 1997. Monte Verde: a Late Pleistocene Am. J. Hum. Genet., 82: 583-592.
settlement in Chile: the archaeological context and Forster P. & Matsumura S. 2005. Did Early
interpretation. Smithsonian Institution Press, Humans Go North or South? Science, 308:
Washington D.C.. 965-966.
Dillehay T.D. 2009. Probing deeper into first Forster P., Harding R., Torroni A. & Bandelt H.J.
American studies. Proc. Natl. Acad. Sci. U.S.A., 1996. Origin and evolution of Native American
106: 971-978. mtDNA variation: a reappraisal. Am. J. Hum.
Dillehay T.D., Ramirez C., Pino M., Collins Genet., 59: 935-945.
M.B., Rossen J. & Pino-Navarro J.D. 2008. Gilbert M.T.P., Jenkins D.L., Gotherstrom
Monte Verde: seaweed, food, medicine and A., Naveran N., Sanchez J.J., Hofreiter M.,
the peopling of South America. Science, 320: Thomsen P.F., Binladen J., Higham T.F.G.,
784-786. Yohe II R.M., Parr R., Cummings L.S. &
Dixon E.J. 2011. Late Pleistocene colonization Willerslev E. 2008a. DNA from pre-Clovis
of North America from Northeast Asia: new human coprolites in Oregon, North America.
insights from large-scale paleogeographic re- Science, 320: 786-789.
constructions. Quat. Int. (in press) (DOI: http:// Gilbert M.T.P., Kivisild T., Gronnow B.,
dx.doi.org/10.1016/j.quaint.2011.02.027). Andersen P.K., Metspalu E., Reidla M., Tamm
Drummond A.J., Rambaut A., Shapiro B. & E., Axelsson E., Gotherstrom A., Campos P.F.,
Pybas O.G. 2005. Bayesan coalescent inference Rasmussen M., Metspalu M., Higham T.F.G.,
and past population dynamics from molecular Schwenninger J.L., Nathan R., De Hoog C.J.,
sequences. Mol. Biol. Evol., 22: 1185-1192. Koch A., Moller L.N., Andreasen C., Meldgaard
Dulik M.C., Zhadanov S.I., Osipova L.P., M., Villems R., Bendixen C. & Willerslev E.
Askapuli A., Gau L., Gokcumen O., Rubistein 2008b. Paleo-Eskimo mtDNA genome reveals
S. & Shurr T.G. 2012. Mitochondrial DNA matrilineal discontinuity in Greenland. Science,
and Y chromosome variation provides evidence 320: 1787-1789.
for a recent common ancestry between Native Goebel T., Waters M.R. & O’Rourke D.H. 2008.
Americans and indigenous Altaians. Am. J. The Late Pleistocene dispersal of modern hu-
Hum. Genet., 90: 229-246. mans in the Americas. Science, 319: 1497-1502.
Endicott P., Gilbert M.T.P., Stringer C.B., Lalueza- Gonzalez-Jose R., Gonzalez-Martin A., Hernandez
Fox C., Willerslev E., Hansen A.J. & Cooper M., Pucciarelli H., Sardi M., Rosales A. & Van
A. 2003. The Genetic Origins of the Andaman Der Molen S. 2003. Craniometric evidence
Islanders. Am. J. Hum. Genet., 72: 178-184. for Palaeoamerican survival in Baja California.
Endicott P., Ho S.Y.W. & Stringer C.B. 2010. Nature, 425: 62-65.
Using genetic evidence to evaluate four pal- Gonzalez-Jose R., Neves W.A., Lahr M.M.,
aeoanthropological hypotheses for the timing Gonzalez S., Pucciarelli H., Martinez M.H. &
of Neanderthal and modern human origins. J. Correal G. 2005. Late Pleistocene/Holocene
Hum. Evol., 59: 87-95. craniofacial morphology in Mesoamerican
Fagan B.M. 2004. The great journey. The peopling Paleoindians: implications for the peopling of
of ancient America. University of Florida Press. the New World. Am. J. Phys. Anthropol., 128:
Gainesville. 772-780.
 J. Hawkset al.
A. Marangoni 93

Gonzalez-Jose R., Bortolini M.C., Santos F.R. & Jelinek A. J. 1971. Early man in the New World:
Bonatto S.L. 2008. The peopling of America: a technological perspective. Arctic Anthropol., 8:
craniofacial shape variation on a continental 15-21.
scale and its interpretation from an interdis- Jota M.S., Lacerda D.R., Sandoval J.R., Viera
ciplinary view. Am. J. Phys. Anthropol., 137: P.R., Santos-Lopes S.S., Bisso-Machado R.,
175-187. Paixao-Cortes R., Revollo S., Paz-Y-Mino C.,
Greenberg J.H., Turner II C.G. & Zegura S.L. Fujita R., Salzano F.M., Bonatto S.L., Bortolini
1986. The settlement of the Americas: a com- M.C., Santos F.R. & The Geneographic
parison of the linguistic, dental and genetic evi- Consortium 2011. A new subhaplogroup of
dence. Curr. Anthropol., 27: 477-497. Native Americans Y-Chromosomes from the
Greyson D.K. & Meltzer D.J. 2003. A requiem Andes. Am. J. Phys. Anthropol., 146: 553-559.
for North American overkill. J. Archeol. Sci., 30: Joyce D.J. 2006. Chronology and new research on
585-593. the Schaefer mammoth (Mammuthus primige-
Guthrie R.D. 2001. Origin and causes of the nius) site, Kenosha County, Wisconsin, U.S.A.
mammoth steppe: a story of cloud cover, wool- Quatern. Int., 142-143: 44-57.
ly mammal tooth pits, buckles, and inside-out Karafet T.M., Zegura S.L., Posukh O., Ospiva
Beringia. Quaternary Sci. Rev., 20: 549-574. L., Bergen A., Long J., Goldman D., Klitz W.,
Harpending H.C., Batzer M.A., Gurven M., Jorde Harihara S., De Knijff P., Wiebe W., Griffith
L.B., Rogers A.R. & Sherry S.T. 1998. Genetic R.C., Templeton A.R. & Hammer M.F.
traces of ancient demography. Proc. Natl. Acad. 1999. Ancestral Asian source(s) of New World
Sci. U.S.A., 95: 1961-1967. Y-chromosome founders haplotypes. Am. J.
Haynes G.A. 2002. The early settlement of North Hum. Genet., 64: 817-831.
America. The Clovis Era. Cambridge University Karafet T.M., Zegura S.L. & Hammer S.F.
Press, Cambridge (U.K.). 2006. Y Chromosomes. In D. Ubelaker (ed):
Howard E.B. 1933. Association of artifacts with Handbook of North American Indians, Vol.
mammoth and bison in eastern New Mexico. 3. Environment, origins and population, pp.
Science, 78: 524. 831-839. Washington D.C., Smithsonian
Hrdlicka A. 1937. The origin and antiquity of the Institution.
American Indian. Annual Report of the Smithsonian Karafet T.M., Mendez F.L., Meilerman M.B.,
Institution, Washington 1923: 481-494. Underhill P.A., Zegura S.L. & Hammer S.F.
Hubbe M., Neves W.A., Atui J.P.V., Cartelle C. & 2008. New binary polimorfisms reshape and
Silva M.P. 2004. A new early human skeleton increase of the human Y Chromosomal haplo-
from Brazil: further support to the ‘‘two main bi- group tree. Genome Res., 18: 830-838.
ological components model’’ for the settlement Kashani B.H., Perego U.A., Olivieri A., Angerhofer
of the Americas. Curr. Res. Pleist., 21: 77-81. N., Gandini F., Carossa V., Lancioni H., Semino
Hubbe M., Neves W.A. & Harvati K. 2010. O., Woodward S.R., Achilli A. & Torroni A.
Testing evolutionary and dispersion scenarios 2011. Mitochondrial haplogroup C4c: a rare
for the settlement of the New World. Plos One, lineage entering America through the ice-free
5: e11105. corridor? Am. J. Phys. Anthropol., 39: 35-39.
Hubbe M., Harvati K. & Neves W.A. 2011. Kemp B.M., Malhi R.S., Mc Donough J., Bolnick
Paleoamerican morphology in the context of D.A., Eshleman J.A., Rickards O., Martinez-
European and east Asian late Pleistocene variation: Labarga C., Johnson J.R., Lorenz J.G., Dixon
implications for human dispersion into the New J., Fifield T.E., Heaton T.H., Worl R. & Smith
World. Am. J. Phys. Anthropol., 144: 442-453. D.G. 2007. Genetic analysis of Early Holocene
Jantz R.L. & Owsley D.W. 2001. Variation skeletal remains from Alaska and its implica-
among Early North American crania. Am. J. tions for the settlement of the Americas. Am. J.
Phys. Anthropol., 114: 146-155. Phys. Anthropol., 132:605–621.

www.isita-org.com
 94 The colonization of the American continent

Kitchen A., Miyamoto M.M. & Mulligan C.J. Mazieres S. 2011. Towards a reconciling model
2008. A three-stage colonization model for the about the initial peploing of America. C. R.
peopling of the Americas. Plos One, 3: e1596. Biologies, 334: 497-504.
Kumar S., Bellis C., Zlojutro M., Melton P.E., McDougall I., Brown F.H. & Fleagle J.G. 2008.
Blangero J. & Curran J.E. 2011. Large scale mi- Sapropels and the age of hominins Omo I and
tochondrial sequencing in Mexican Americans II, Kibish, Ethiopia. J. Hum. Evol., 55: 409-420.
suggests a reappraisal of Native American ori- Mellars P. 2006. Going East: New genetic and ar-
gins. BMC Evol. Biol., 11: 293. chaeological perspectives on the modern human
Lahr M.M. 1995. Patterns of modern human di- colonization of Eurasia. Science, 313: 796-800.
versification: implications for Amerindian ori- Meltzer D.J. 2009. First peoples in a New World.
gins. Yearb. Phys. Anthropol., 38: 163-198. Colonizing Ice Age America. University of
Lyman R.L., O’Brien M. & Hayes V. 1998. A me- California Press. London.
chanical and functional study of bone rods from Mulligan C.J., Hunley K., Cole S. & Long J.C.
the Richey-Roberts Clovis cache, Washington, 2004. Population genetics, history and health
USA. J. Archeol. Sci., 25: 887-906. patterns in Native Americans. Annu. Rev.
Macaulay V., Hill C., Achilli A., Rengo C., Clarke Genomics Hum. Genet., 5: 295-315.
D., Meehan W., Blackburn J., Semino O., Scozzari Mulligan C.J., Kitchen A. & Miyamoto M.M.
R., Cruciani F., Taha A., Shaari N.K., Raja J.M., 2008. Updated Three-Stage model for the peo-
Ismail P., Zainuddin Z., Goodwin W., Bulbeck pling of the Americas. Plos One, 3: e3199.
D., Bandelt H.J., Oppenheimer S., Torroni A. & Neves W.A. & Hubbe M. 2005. Cranial mor-
Richards M. 2005. Single, rapid coastal settlement phology of early Americans from Lagoa Santa,
of Asia revealed by analysis of complete mitochon- Brazil: implications for the settlement of the
drial genomes. Science, 308: 1034-1036. New World. Proc. Natl. Acad. Sci. U.S.A., 102:
Malhi R.S., Kemp B.M., Eshleman J.A., Cybulski 18309-18314.
J., Smith D.G., Cousins S. & Harry H. 2007. Neves W.A & Pucciarelli H. 1991. Morphological
Mitochondrial haplogroup M discovered in affinities of the first Americans: an exploratory
prehistoric North Americans. J. Archeol. Sci., analysis based on early South American human
34:642-648. remains. J. Hum. Evol., 21: 261-273.
Mandryk C.A.S., Josenhans H., Fedje D.W. Neves W.A. & Pucciarelli H. 1998. The
& Mathewes R.W. 2001. Late Quaternary Zhoukoudian Upper Cave skull 101 as seen
paleoenvironments of Northwestern North from the Americas. J. Hum. Evol., 34: 219-222.
America: implications for inland versus coast- Neves W.A., Powell J.F. & Ozolins E.G. 1999.
al migration routes. Quaternary Sci. Rev., 20: Extra-continental morphological affinities of
301-314. Palli Aike, southern Chile. Interciencia, 24:
Manzi G. 2012. On the trail of the genus Homo 258-263.
between archaic and derived morphologies. J. Neves A.W., Hubbe M., Mercedes M., Okumura
Anthropol. Sci., 90: 99-116. M., Gonzalez-Jose R., Figuti L., Eggers S. & De
Manzi G., Grimaldi S. & Destro-Bisol G. 1994. Blasis P.A.D. 2005. A new early Holocene hu-
Modern human origin/s: fossil, genetic and ar- man skeleton from Brazil: implications for the
cheological evidence. State of the art. Rivista di settlement of the New World. J. Hum. Evol.,
Antropologia, 2: 243-269. 48: 403-414.
Martin P.S. 1973. The discovery of America. O’Fallon B.D. & Fehren-Schmitz L. 2011. Native
Science, 179: 969-974. Americans experienced a strong population bot-
Martin P.S. 1984. Prehistoric overkill: The global tleneck coincident with European contact. Proc.
model. In P.S. Martin & R.G. Klein (eds): Natl. Acad. Sci. U.S.A., 108: 20444-20448.
Quaternary extinctions: A prehistoric revolution, O’Rourke D.H. 2006. Blood groups, immuno-
pp 354-403. University of Arizona Press, Tucson. globulins and genetic variation. In D. Ubelaker
 J. Hawkset al.
A. Marangoni 95

(ed): Handbook of North American Indians. Vol. and evolutionary diversification in America.
3. Environments, origins and population, pp. 762- Quatern. Int., 109-110: 123-132.
776. Smithsonian Institution, Washington D.C.. Raff J.A., Bolnick D.A., Tackney J. & O’Rourke
O’Rourke D.H. & Raff J.A. 2010. The human D.H. 2011. Ancient DNA perspective on
genetic history of the Americas: the final fron- American colonization and population hystory.
tier. Curr. Biol., 20: 202-207. Am. J. Phys. Anthropol., 146: 503-514.
Peltier W.R. 2002. On eustatic sea level his- Rasmussen M., Li Y., Lindgreen S., Pedersen
tory: Last Glacial Maximum to Holocene. J.S., Albrechtsen A., Moltke I., Metspalu M.,
Quaternary Sci. Rev., 24: 377-396. Metspalu E., Kivisild T., Gupta R., Bertalan
Perego U.A., Achilli A., Angerhofer N., Accetturo M., Nielsen K., Gilbert M.T., Wang Y.,
M., Pala M., Olivieri A., Kashani B.H., Ritchie Raghavan M., Campos P.F., Kamp H.M.,
K.H., Scozzari R., Kong Q.P., Myres N.M., Wilson A.S., Gledhill A., Tridico S., Bunce M.,
Salas A., Semino O., Bandelt H.J., Woodward Lorenzen E.D., Binladen J., Guo X., Zhao J.,
S.R. & Torroni A. 2009. Distinctive but con- Zhang X., Zhang H., Li Z., Chen M., Orlando
comitant Paleo-Indian migration routes from L., Kristiansen K., Bak M., Tommerup
Beringia marked by two rare mtDNA haplo- N., Bendixen C., Pierre T.L., Gronnow B.,
groups. Curr. Biol., 19: 1-8. Meldgaard M., Andreasen C., Fedorova S.A.,
Perego U.A., Angerhofer N., Pala M., Olivieri A., Osipova L.P., Higham T.F., Ramsey C.B.,
Lancioni H., Kashani B.H., Carossa V., Ekins J.E., Hansen T.V., Nielsen F.C., Crawford M.H.,
Gomez-Carballa A., Huber G., Zimmermann B., Brunak S., Sicheritz-Ponten T., Villems R.,
Corach D., Babudri N., Panara F., Myres N.M., Nielsen R., Krogh A., Wang J. & Willerslev E.
Parson W., Semino O., Salas A., Woodward S.R., 2010. Ancient human genome sequence of an
Achilli A. & Torroni A. 2010. The initial peopling extinct Palaeo-Eskimo. Nature, 463: 757-762.
of the Americas: a growing number of founding Ray N., Wegmann D., Fagundes N.J.R., Wang S.,
mitochondrial genomes from Beringia. Genome Ruiz-Linares A. & Excoffier L. 2009. A statis-
Res., 20: 1174-1179. tical evaluation of models for the initial settle-
Perez S.I., Bernal V., Gonzalez P.N., Sardi M. & ment of the American continent emphasizes the
Politis G.G. 2009. Discrepancy between cranial importance of gene flow with Asia. Mol. Biol.
and DNA data of early Americans: implications Evol., 27: 337-345.
for American peopling. Plos One, 4: e5746. Reich D., Patterson N., Campbell D., Tandon
Pitulko V.V., Nikolsky P.A., Girya E.Y., Basilyan A., Mazieres S., Ray N., Parra M.V., Rojas W.,
A.E., Tumskoy V.E., Koulakov S.A., Astakhov Duque C., Mesa N., Garcıa L.F., Triana O., Blair
S.N., Pavlova E.Y & Anisimov M.A. 2004. The S., Maestre A., Dib J.C., Bravi C.M., Bailliet
Yana RHS: humans in the artic before the Last G., Corach D., Hunemeier T., Bortolini M.C.,
Glacial Maximum. Science, 303: 52-56. Salzano F.M., Petzl-Erler M.L., Acuna-Alonzo
Powell J.F. & Neves W.A. 1999. Craniofacial mor- V., Aguilar-Salinas C., Canizales-Quinteros S.,
phology of the first Americans: pattern and pro- Tusie-Luna T., Riba L., Rodrıguez-Cruz M.,
cess in the peopling of the New World. Yearb. Lopez-Alarcon M., Coral-Vazquez R., Canto-
Phys. Anthropol., 42: 153-188. Cetina T., Silva-Zolezzi I., Fernandez-Lopez
Powell J.F., Neves W.A., Ozolins E. & Pucciarelli J.C., Alejandra V. Contreras A.V., Jimenez-
H.M. 1999. Afinidades biologicas extra-con- Sanchez G., Gomez-Vazquez M.J., Molina J.,
tinentales de lo dos esqueletos mas antiguos Carracedo A., Salas A., Gallo C., Poletti G.,
de America: implicaciones para el poblami- Witonsky D.B., Alkorta-Aranburu G., Sukernik
ento del Nuevo Mundo. Antropologia Física R.I., Osipova L., Fedorova S.A., Vasquez R.,
Latinoamericana, 2: 7:22. Villena M., Moreau C., Barrantes R., Pauls
Pucciarelli H., Sardi M.L., Jimenez Lopez J.C. D., Excoffier L., Bedoya G., Rothhammer F.,
& Serrano Sanchez C. 2003. Early peopling Dugoujon J.M., Larrouy G., Klitz W., Labuda

www.isita-org.com
 96 The colonization of the American continent

D., Kidd J., Kidd K., Di Rienzo A., Freimer populations. Yearb. Phys. Anthropol. 24:
N.B., Price A.L. & Ruiz-Linares A. 2012. 37–73.
Reconstructing Native American population Szathmáry E.J.E. 1984. Peopling of Northern
history. Nature, 488: 370–374. North America: Clues from Genetic Studies.
Rose J.I., Usik V.I., Marks A.E., Hilbert Y.H., Acta Anthropogenetica, 8: 79-109.
Galletti C.S., Parton A., Geiling J.M., Cerny Szathmáry E.J.E. 1996. Ancient Migrations from
V., Morley M.W., & Roberts R.G. 2011. The Asia to North America. In T. Akazawa, &
Nubian Complex of Dhofar, Oman: an African E.J.E. Szathmary (eds): Prehistoric Mongoloid
Middle Stone Age Industry in Southern Arabia. Dispersals, pp 149–164. Oxford Univ. Press,
Plos One, 4: e28239. Oxford.
Rothhammer F. & Dillehay T.D. 2009. The Late Tamm E., Kivisild T., Reidla M., Metspalu
Pleistocene Colonization of South America: M., Smith D.G., Mulligan C.J., Bravi,
an Interdisciplinary Perspective. Ann. Hum. C.M., Rickards O., Martinez-Labarga
Genet., 73: 540-549. C., Khusnutdinova E.K., Fedorova., S.A.,
Scabuzzo C. & Politis G. 2007. Early-Holocene Golubenko M.V., Stepanov V.A., Gubina
secundary burials in the Pampas of Argentina. M.A. Zhadanov S.I., Ossipova L.P., Damba
Curr. Res. Pleist., 23: 64-66. L., Voevoda M.I., Dipierri J.E., Villems R.
Schroeder K.B., Jakobsson M., Crawford M.H., & Malhi R.S. 2007. Beringian standstill and
Schurr T.G., Boca S.M., Conrad D.F., Tito spread of Native American founders. Plos One,
R.Y., Osipova L.P., Tarskaia L.A., Zhadanov 3: e829.
S.I., Wall J.D., Pritchard J.K., Malhi R.S., Smith Taylor R.E., Kirner D.L., Southon J.R. & Chatters
D.G. & Rosenberg N.A. 2009. Haplotypic J.C. 1998. Radiocarbon Dates of Kennewick
background of a private allele at high frequency Man. Science, 280: 1171.
in the Americas. Mol. Biol. Evol., 26: 995-1016. Thangaraj K., Chaubey G., Kivisild T., Reddy
Schurr T.G. & Sherry S.T. 2004. Mitochondrial A.G., Singh V.K., Rasalkar A.A. & Sing L.
DNA and Y chromosome diversity and the 2005. Reconstructing the Origin of Andaman
peopling of the Americas: evolutionary and Islanders. Science, 308: 996.
demographic evidence. Am. J. Hum. Biol., 16: Torroni A., Schurr T.G., Yang C.C., Szathmary
420-439. E.J.E., Williams R.C., Schanfield M.S., Troup
Schurr T.G., Ballinger S.W., Gan Y.Y., Hodge G.A., Knowler W.C., Lawrence D.N., Weisss
J.A., Merriwether D.A., Lawrence D.N., K.M. & Wallace D.C. 1992. Native American
Knowler W.C., Weiss K.M. & Wallace D.C. mitochondrial DNA analysis indicates that
1990. Amerindian mitochondrial DNAs have the Amerind and the Nadene populations
rare Asian mutations at high frequencies, sug- were founded by two independent migrations.
gesting they derived from four primary mater- Genetics, 130: 153-162.
nal lineages. Am. J. Hum. Genet., 46: 613-623. Torroni A., Schurr T.G., Cabell M.F., Brown
Straus L.G., Meltzer D.J. & Goebel T. 2005. Ice M.D., Neel J.V., Larsen M., Smith D.G., Vullo
Age Atlantis? Exploring the Solutrean-Clovis C.M. & Wallace D.C. 1993. Asian affinities
“Connection”. World Archaeol., 37: 507-532. and the continental radiation of the four found-
Stringer C.B. 2002. Modern human origins: pro- ing Native American mtDNAs. Am. J. Hum.
gress and prospects. Proc. R. Soc. Lond. B. Sci., Genet., 53: 563-590.
357: 563-579. Van Vark G.N., Kuizenga D. & Williams F. L’E.
Stringer C.B. & Andrews P. 1988. Genetic and 2003. Kennewick and Luzia: lessons from
fossil evidence for the origin of modern hu- the European Upper Paleolithic. Am. J. Phys.
mans. Science, 239: 1263-1268. Anthropol., 121: 181-184.
Szathmáry E.J.E. 1981. Genetic markers in Wang S., Lewis Jr. C.M., Jakobsson M.,
Siberian and northern North American Ramachandran S., Ray N., Bedoya G., Rojas W.,
 J. Hawkset al.
A. Marangoni 97

Parra M.V., Molina J.A., Gallo C., Mazzotti G., Webb S.D. 2005. First Floridians and Last
Poletti G., Hill K., Hurtado A.M., Labuda D., Mastodons. The Page-Ladson site in the Aucilla
Klitz W., Barrantes R., Bortolini M.C., Salzano River. Springer, Dordrecht, The Netherlands.
F.M., Petzl-Erler M.L., Tsuneto L.T., Llop E., Williams R.C., Steinberg A.G., Gershowitz H.,
Rothhammer F., Excoffier L., Feldman M.W., Bennett P.H., Knowler W.C., Pettitt D.J., Butler
Rosenberg N.A. & Ruiz-Linares A. 2007. W., Baird R., Dowda-Rea L. & Burch T.A. 1985.
Genetic variation and population structure in GM allotypes in Native Americans: evidence for
Native Americans. Plos Genet., 3: 2049-2067. three distinct migrations across the Bering land
Waters M.R. & Stafford Jr. T.W. 2007. Redefining bridge. Am. J. Phys. Anthropol., 66: 1-19.
the age of Clovis: implications for the peopling Zegura S.L., Karafet T.M., Zhivotovsky L.A. &
of the Americas. Science, 315: 1122-1126. Hammer M.F. 2004. High-Resolution SNPs
Waters M.R., Forman S.L., Jennings T.A., Nordt and microsatellite haplotypes point to a single,
L.C., Driese S.G., Feinberg J.M., Keene J.L., recent entry of Native American Y chromo-
Halligan J., Lindquist A., Pierson J., Hallmark somes into the Americas. Mol. Biol. Evol., 21:
C.T., Collins M.B. & Wiederhold J.E. 2011. 164-175.
The Buttermilk Creek Complex and the
Origins of Clovis at the Debra L. Friedkin Site,
Texas. Science, 331: 1599-1603. Editor, Giovanni Destro-Bisol

www.isita-org.com