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Genetic Heterogeneitybetween Berbers andArabsLara R Arauna,
Institut de Biologia Evolutiva (CSIC-UPF), Departament de
CiènciesExperimentals i de la Salut, Universitat Pompeu Fabra,
Barcelona, Spain
David Comas, Institut de Biologia Evolutiva (CSIC-UPF),
Departament de CiènciesExperimentals i de la Salut, Universitat
Pompeu Fabra, Barcelona, Spain
Advanced article
Article Contents• Introduction
• Genetic Diversity in Berber and Arab Groups
• Related Articles
Online posting date: 15th September 2017
The human population history of North Africahas been different
from the rest of the conti-nent, and it has been characterised by
populationreplacements, extensive continuous gene flow,and
differential admixture from neighbouringregions. This complex
demographic landscape hasyielded a large degree of genetic
heterogeneityamong North African populations. Recent histori-cal
admixture processes have been inferred fromgenome-wide data; no
correlation between genet-ics and ethnic groups has been described,
pointingto a lack of genetic differentiation between Berberand Arab
groups in North Africa. This complexgenetic population structure
should be taken intoaccount when designing biomedical approaches
inNorth African groups.
Introduction
Little is known about the initial human settlement in
NorthAfrica, the movements during the Upper Paleolithic (known
asthe ‘Late Stone Age’ in the study of African prehistory) andthe
possible introgression from archaic humans, such as Nean-derthals
(Green et al., 2010) and Denisovans (Reich et al., 2010)or other
unknown African hominins (Hammer et al., 2011), tothe populations
inhabiting this area. Recent description of formsof Homo sapiens
with a mosaic of modern and primitive char-acteristics in North
Africa dated back to ∼300 000 ya (yearsago) has even challenged the
geographical origins of our species(Hublin et al., 2017). However,
the first presence of anatomi-cally modern humans (AMHs) in North
Africa has been estimated190 000–130 000 ya by archaeological
studies (much older thanprevious estimates that dated the first
occupation of the region
eLS subject area: Evolution and Diversity of Life
How to cite:Arauna, Lara R and Comas, David (September 2017)
GeneticHeterogeneity between Berbers and Arabs. In: eLS. John
Wiley& Sons, Ltd: Chichester.DOI:
10.1002/9780470015902.a0027485
∼45 000 ya) (Smith et al., 2007). The Aterian is the first
prehis-toric industry characterised in North Africa (Barton et al.,
2009),which has been suggested to have started around 120 000
ya.Although paradoxical, the Sahara desert has been reported to bea
corridor for the movement of people during the ‘green
Sahara’periods thanks to its watercourses. The old dates of AMH
remainsand the Sahara corridor leave many open questions about the
roleof North Africa in the origin of modern humans and their
disper-sal out of the continent. Different hypotheses have been
proposed:Was North Africa just a stop on the road out of Africa, or
did itplay a more important role in the evolution of modern
humans?
No clear connections have been established between this
firsthuman industry and subsequent cultures in the region, such as
theIberomaurusian industry (22 000–9500 ya) (Newman, 1995).
Theprefix ‘Ibero-’ refers to the ancient presumption that this
cultureextended into Iberia, although an origin in the Nile River
valley iswidely accepted (Camps, 1995). The Iberomaurusian culture
wasfollowed by the Capsian industry (10 000–4700 ya) (Newman,1995)
that persisted well into the Neolithic, which began around5500 ya
in the region. The prehistoric cultural changes in NorthAfrica were
independent of the change dynamics on the Europeanshores of the
Mediterranean.
Based on archaeology and linguistics, it has been arguedthat the
peopling of North Africa has not been continuous andthat population
replacements may have occurred in differentmoments. Genetic data
(Henn et al., 2012) has supported thispopulation replacement
hypothesis in North Africa, suggesting aback-to-Africa migration
from the Middle East in pre-Holocenetimes, older than 12 000 ya.
The ancestors of today’s NorthAfrican populations derive at least
partially from this migrationwave; however, it is not clear if this
replacement was completeor there might be some traces of ancient
continuity in the region.Therefore, many open questions remain
about the demography ofthe region in relation to the demographic
continuity of humanssince the first occupation of North Africa by
modern humans,such as how many times a population replacement took
place andwhich are the demographic parameters such as population
size ofthe migrants and where did they come from.
The challenges of the understanding of North African
pop-ulations are not limited to prehistoric times: in historic
times,North Africa has also experienced a very complex history that
isreflected in its human demography. Historical records
documenttrade routes across the Sahara and contacts between both
Mediter-ranean shores and the Middle East. Phoenicians (814 B.C.)
and
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Genetic Heterogeneity between Berbers and Arabs
Romans (146 B.C.) occupied part of coastal North Africa
withlimited population contributions, which were even less
signifi-cant for Vandals (A.D. 429) and Byzantines (A.D. 533)
(New-man, 1995). One of the major historical events in the
regionthat changed the cultural and demographic landscape of
NorthAfrica is the Arabisation. The first Arab invasion, initially
con-fined to Egypt, started in A.D. 643 and may have involved only
afew thousand individuals (McEvedy, 1995). The Arabs began toimpose
their religion and language over the Berber
autochthonouspopulation, a process that culminated with a second
and largerArab wave in which the Bedouin reached the Maghreb
(northwestAfrica) in the eleventh century. The Islamic expansion
even wenton to engulf the Iberian Peninsula (A.D. 711). The later
arrivalsto North Africa in colonial times include Europeans and
OttomanTurks, mainly in Egypt.
Nowadays in North Africa, two main groups of languages
arespoken: Berber and Arab dialects. Both belong to the
Afroasiaticfamily, which is a group of languages that might have
originatedin the Mesolithic, between 15 000 and 10 000 ya. Although
it isnot clear, one of the most plausible places of origin of
Afroasiaticlanguages is the ‘grassland east of the Nile Valley’
(Newman,1995). However, it is unknown if the arrival of the
Afroasiaticlanguages was mainly a cultural diffusion process or it
alsoimplied a relevant demographic movement.
The Berbers (Amazighen, as they call themselves) are con-sidered
the autochthonous people of North Africa. As can beelucidated from
the complex and unsolved prehistory and his-tory of North Africa,
it is difficult to establish an origin for theBerbers, and many
hypotheses have been proposed. Historicalrecords reveal their
presence before the Phoenician arrival tothe region (Camps, 1998),
and archaeologically, it has been sug-gested a link between the
Capsian culture and the Berber people(Camps, 1995). However, the
Berbers might not have been a uni-form and homogeneous group, since
it has been known that thepopulations that inhabited North Africa
by the time of the Phoeni-cian arrival were a composite group known
as Libyco-Berbers,where we find the Garamantes, Bavares, Mauri,
Gaetuli, amongmany others (Newman, 1995). In addition to the
uncertaintiesabout the origins of the Berbers, the arrival of other
people inthe region, especially the influence of Arabs, makes the
under-standing of the population history of Berbers more
challenging.
Considering these previous points, the population genetics
def-inition of today’s Berber and Arab groups is a complex
task.Different points of view can be considered to determine
whethera population is considered as Berber or not. Berbers can be
under-stood as non-Semitic autochthonous populations in North
Africa,although it is challenging to determine which current
populationsderive from an ancestral autochthonous group in North
Africa.Another way to identify Berbers and Arabs is regarding their
cul-tural characteristics, but again, the cultural exchange in
recenthistory of North Africa has been common, and it is difficult
todifferentiate, for example, an Arabised Berber society from
anArab population in North Africa, among many other combina-tions.
Given this combination of circumstances, the best optionfor a
classification may be based on linguistics, Berber popula-tions
being those who speak Berber languages and Arabs beingthose
speaking Arab (Dugoujon et al., 2009). This classifica-tion could
be the way of having less ‘false positives’; however,
this classification is not free of caveats, and populations
identi-fied as Arabs or non-Berbers could in fact have a large
Berbergenetic ancestry. It is important to emphasise that the
purpose ofthis classification is to understand the genetics of
North Africanpopulations and in any case to establish
socio-cultural groups,which should take into consideration many
other aspects (tradi-tions, identity, etc.).
Despite the scanty genetic data on North African groups,
pop-ulation genetics might shed some light on the genetic structure
ofBerbers and Arabs, trying to establish the demographic
scenariosthat have modelled the current genetic landscape in North
Africa.
Genetic Diversity in Berber andArab Groups
The human genetic data of North Africa is scarce compared
toother regions of the African continent, and most of the
Africangenetic diversity studies have been focused on the origin of
ourspecies and the first dispersions out of Africa (see, for
instance,Tishkoff et al., 2009), the genetic diversity of North
Africanpopulations and surrounding groups being poorly
characterised.
The analyses based on frequencies of classical genetic
poly-morphisms (blood groups, red cell enzymes and serum
proteins)have shown that the genetic landscape in North Africa
presents aneast-west pattern of variation without differences
between Arabsand Berbers, pointing to a sizeable Upper Paleolithic
componentin current North African populations, whereas the
Neolithic dif-fusion in the region was more a cultural than a demic
process(Barbujani et al., 1994; Bosch et al., 1997). These
classical stud-ies showed little contribution from sub-Saharan
populations inNorth Africans and genetic differentiation from
southern Euro-peans, including Iberians (Bosch et al., 1997).
However, the anal-ysis of mtDNA lineages has shown an important
sub-Saharancontribution, although most haplogroups in North Africa
are ofwest-Eurasian origin (Fadhlaoui-Zid et al., 2004, 2011b;
Kringset al., 1999; Plaza et al., 2003; Rando et al., 1998). Some
ofthem can be traced to ancient Paleolithic times (such as
hap-logroups U6, M1, which are almost specific of North
Africanpopulations); however, some maternal lineages have been
theresult of a more recent acquisition from Europe or the Mid-dle
East (such as haplogroups U5, V, R0a, J1b, U3) (Gonzálezet al.,
2007; Maca-Meyer et al., 2003; Olivieri et al., 2006)(Figure 1). In
addition, a large degree of genetic heterogene-ity has been shown
in North African maternal and paternal lin-eages compared to other
geographical regions such as Europe(Fadhlaoui-Zid et al., 2004,
2011a; Plaza et al., 2003). The anal-ysis of Y-chromosome lineages
has shown a high frequency oftwo specific North African haplogroups
(E-M81 and E-M78)(Figure 1), although their origins have been
controversial sincesome analyses have suggested a Paleolithic
component (Boschet al., 2001), whereas others have pointed to a
Neolithic ori-gin (Arredi et al., 2004; Cruciani et al., 2004,
2007; Seminoet al., 2004). The data on autosomal markers in North
Africanpopulations based on some short tandem repeats (STRs)
(Boschet al., 2000) and Alu polymorphisms (Comas et al., 2000;
Flo-res et al., 2000; González-Pérez et al., 2010) has been
analysed
2 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
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Genetic Heterogeneity between Berbers and Arabs
Siwa
M
ozab
ite
Sen
edM
atm
ata
Chen
ini D
ouire
t
Souss
Figuig
Bouhria
Ansi
Tu
nisian
s
Algerians
Saharawis
Moroccans
Mt DNA haplogroups
Ychr haplogroups
A3b2*
E3a*
E1
E3b3a*
E3b3*
E3b1*
E3b*
E3b2a
E3b2*
E3b2b
I1b2*
I*(xI1a2.I1a3.I1b2)
J2f1
J2*
J2a
F*(xH.I.J.K)
J*
O*(xO1.O2a.O3)
K*(xK2.K3.L.M.N1.N3.O.P)
K2
R1b8
R1*(xR1a1.R1b1-R1b8)
R1a1*
P*(xP1.Q1.Q2.Q3.R1)
J2f*
L0
L1
L2
L3
L4NAL5
M1
M
U*
U6
K
HV
H
HV0
Other
N1/N2
I
X
J/T
T
J
Arabs
Berbers
MoroccoAlgera
TunisiaEgypt
ArabsBerbers
MoroccoAlgeraTunisiaEgypt
100
75
50
25
0
Fre
q
100
75
50
25
0
Fre
q
Algeria
ns
Sou
th M
oroc
cans
Nor
th M
oro
ccan
s
South
Egyp
tians
North Egyptians
Tunisians
Alg
erians
Saharawis
Moroccans
(a)
(b)
Figure 1 Haplogroup frequencies for different Berber and Arab
populations for (a) mitochondrial DNA (data from Coudray et al.,
2009 and Plaza et al.,2003) and (b) Y chromosome (data from Arredi
et al., 2004).
eLS © 2017, John Wiley & Sons, Ltd. www.els.net 3
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Genetic Heterogeneity between Berbers and Arabs
in few North African samples pointing to a closer genetic
affin-ity to West Eurasian populations. Y-chromosome lineages
andautosomal markers show a clear differentiation of North
Africanpopulations with some sub-Saharan gene flow and almost
negli-gible European contact (Bosch et al., 2000, 2001; Comas et
al.,2000; González-Pérez et al., 2010). Nonetheless, the
contributionof North African female and male lineages in the
Iberian Penin-sula (Adams et al., 2008; Plaza et al., 2003) should
be stressed.
Genome-wide analyses have recently become the most pow-erful
tool to unravel the genetic history of populations. The firstlarge
genome-wide study in North Africa analysing several NorthAfrican
and surrounding populations (Henn et al., 2012) sug-gested the
existence of an autochthonous North African compo-nent whose origin
is located out of Africa before the Holocene(>12 000 years ago)
and a complex admixture from the Mid-dle East, Europe and
sub-Saharan Africa. This autochthonousNorth African component was
found in high frequencies in theonly Berber sample (Berber Tunisian
from Chenini) analysed inthe study, suggesting that Berbers might
be responsible for thisautochthonous component in North African
samples. However,recent genome-wide data with additional Berber
samples haveshown a different scenario, with some Berber groups
present-ing less frequencies of the autochthonous component
comparedto some Arab groups (Arauna et al., 2017). In addition to
theautochthonous component in North Africa, it has been possible
toquantify and estimate the recent dates of the sub-Saharan
admix-ture in North Africa (Henn et al., 2012) as well as the
geneticcontacts from North Africa mainly restricted to
south-westernEurope (Botigué et al., 2013). Concerning the
introgression ofarchaic hominins into North Africans, some
Neanderthal admix-ture is detected in these groups independent of
their Europeancomponent (Sánchez-Quinto et al., 2012).
Most of the genetic studies in North African populations
agreewith a limited or non-existent correlation between genetics
andgeography, and therefore, they show a high population
hetero-geneity in the region (Bosch et al., 2000; Fadhlaoui-Zid et
al.,2004; Flores et al., 2011; González-Pérez et al., 2010).
How-ever, an east-west genetic pattern of variation has been
observed(Arauna et al., 2017; Bosch et al., 1997; Harich et al.,
2001; Hennet al., 2012), which has been explained by Middle Eastern
geneflow towards North Africa according to genome-wide
analyses(Arauna et al., 2017; Henn et al., 2012). The genetic
heterogene-ity is not only due to the wide extension of North
Africa, since itis also found at microgeographical level when
considering morerestricted areas, such as Algeria (Bekada et al.,
2015) or Tunisia(Fadhlaoui-Zid et al., 2011a; Kefi et al., 2015),
for instance.Regardless of the genetic markers analysed, North
African popu-lations have been described as a mosaic of North
African, MiddleEastern, European and sub-Saharan ancestries. The
differentialadmixture with these four ancestry sources explains the
currentgenetic structure in North Africa, characterised by diverse
andheterogeneous populations. This differential admixture
explainswhy nearby populations or even individuals inhabiting the
samelocation might be genetically more distant than groups of
peo-ple in geographically distant populations. For instance,
NorthAfrican individuals with similar amounts of sub-Saharan
admix-ture present high genetic similarities regardless of their
geograph-ical origin. Neither geography, linguistics nor ethnic
affiliation
is correlated with a specific pattern of admixture proportions.
Acompletely different scenario was shown when analysing
NorthAfrican Jews, who are genetically closer to other Jewish
popula-tions than to their geographic neighbours (Campbell et al.,
2012).There are many possible scenarios that might explain the
dif-ferential admixture in North Africa, socio-cultural structure
andhistorical events being the main reasons. However, more com-plex
socio-cultural scenarios, including socio-economic factors,a more
precise ethnic or linguistic definition, particularities ofthe
history of each local area and rural versus urban lifestyles,should
be considered in order to find a correlation with the com-plex
genetic pattern observed (Arauna et al., 2017).
Concerning the genetic differences between Berber and
Arabgroups, classical and uniparental markers have highlighted
thelack of a genetic structure that could differentiate Berber
andArab populations (Bekada et al., 2015; Fadhlaoui-Zid et al.,
2004;Harich et al., 2001; Kefi et al., 2015). Most of these authors
haveagreed on a cultural rather than a demographic impact of
theArabisation after the Arab expansion (Bosch et al., 2000,
2001;Ennafaa et al., 2011; Flores et al., 2011). However, as stated
ear-lier, an East-to-West genetic gradient of a Middle Eastern
compo-nent has been described in genome-wide analyses, and
admixtureevents have been dated to the Arabisation period (Arauna
et al.,2017; Henn et al., 2012), suggesting that the Arabisation
mighthave had more demographic impact than previously thought.The
first widespread genome-wide study in North Africa (Hennet al.,
2012) suggested that the single Berber sample analysed(Tunisian
Berber from Chenini) presented high frequencies of anautochthonous
North African component (named in the study the‘Maghrebi’
component) and negligible presence of the MiddleEastern component.
This lack of Middle Eastern component inBerbers and its higher
frequencies in Arabs pointed to a genomedifferentiation between
both North African groups. However, theposterior genome-wide
analysis of additional Berber samples haschallenged this genetic
differentiation idea (Arauna et al., 2017).This study has
corroborated the lack of genetic differentiationbetween Berbers and
Arabs; however, it has shown that the Arabi-sation had also a
demographic impact in North Africa (Figure 2).It is difficult to
exactly quantify the magnitude of the Arab geneticcontribution
during the Arab expansion in the seventh century,among other
reasons because the influence from Middle East hasbeen continuous
into North Africa (Camps, 1995). Therefore, thisdata supports the
lack of significant differences between Berberand Arabs, but points
to a new explanation: a high and differentiallevel of admixture
since the Arab expansion that has affected mostNorth African groups
regardless of their Berber or Arab identity.
The other genetic components that are present in North
Africanpopulations also contribute to the heterogeneity between
Berberand Arabs. Both peoples have admixed with sub-Saharan
popula-tions in the last two millennia regardless of their
ethnicity, whichhas contributed to the lack of differentiation
between both groups(Flores et al., 2000; Harich et al., 2010; Henn
et al., 2012). How-ever, it has been shown that the sub-Saharan
admixture is mostfrequent in the southern groups, and in fact, a
South-North gradi-ent of sub-Saharan admixture has been described
in North Africa(Comas et al., 2000; Plaza et al., 2003), which
sometimes canmimic differences between Arab and Berber populations
that are
4 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
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Genetic Heterogeneity between Berbers and Arabs
Morocco S. Morocco N. Sahara Algeria
Algeria Tim.Morocco Err.Morocco TiznitTunisia Sened
Libya
Ancestry
Tun. Chen. Ber.
YRI
Arabs
Berbers
Syria
Basque
Egypt
Figure 2 Analyses of autosomal markers. Each bar shows the
proportion of the genome that each individual share with each of
the four ancestral populationsaccording to ChromoPainter analyses
(Lawson et al., 2012). The separation between bars within each
population represents clusters of individuals with similarancestry
proportions. Source: Arauna et al. (2017).
mainly explained by a differential sub-Saharan contribution
(Flo-res et al., 2011).
The high heterogeneity found in North African populationsis also
present within ethnic groups. Berbers are a compositeof groups with
very different demographic characteristics, andsome Berber groups
are genetic outliers compared to other Berberor North African
populations. This is the case of the TunisianBerber population from
Chenini, which has been analysed foruniparental and genome-wide
markers, and in both cases, it hasbeen described as an outlier
group compared to their NorthAfrican populations (Ennafaa et al.,
2011; Fadhlaoui-Zid et al.,2004, 2011a; Henn et al., 2012). This
genetic singularity isnot exclusive of this group since other
Tunisian Berber groupshave also shown similar outlier
characteristics (Ennafaa et al.,2011; Fadhlaoui-Zid et al., 2004,
2011a). Another example arethe Mozabite, an Algerian Berber group
included in the HumanGenome Diversity Panel (HGDP) (Cann et al.,
2002), whichhave been extensively analysed in many genetic studies
and havealso been described as an outlier group within North Africa
andalso within Berbers (Bekada et al., 2015; Plaza et al., 2003).It
is important to note that Mozabite have been considered asthe
representative North African group in a large number ofstudies due
to the fact that they are the only North Africangroup in the HGDP,
which should be taken with caution giventheir genetic outlier
characteristics. Furthermore, the Siwa Berberpopulation from Egypt
is genetically distant from other Berbergroups from West North
Africa (Coudray et al., 2009), suggestinga differentiation between
Egyptian Berbers and Western Berbers.However, there are many other
Berber groups that are closelyrelated between them and to other
North African groups. All theseevidences support a high diversity
within Berbers, which couldbe explained by isolation processes and
inbreeding. For example,the Chenini population is an isolated
population that shows highlevels of inbreeding (Arauna et al.,
2017; Henn et al., 2012). It hasbeen described that some of the
diversity observed within Berbersis due to the belonging to an
urban or rural area. Rural areas favourisolation leading to genetic
differentiated populations (Brakezet al., 2001; Ennafaa et al.,
2011; Frigi et al., 2006). On theother hand, urban areas that are
less isolated are more favourableto high admixture levels and lead
to the lack of differentiationbetween Berber and Arab groups.
Finally, it is important to
notice the wide extension and fragmentation of the territory
wherethe Berbers live, which could lead to population
differentiationbecause of many reasons apart from ethnicity (simply
by isolationby distance or differential gene flow). Then, it is
important tocompare the Berber groups to geographically close
populationsin order to make inferences of genetic distances.
The genetic heterogeneity of North African samples, the com-plex
population admixture patterns and the lack of correla-tion between
genetics and ethnic groups should be taken intoaccount when
addressing biomedical approaches in North Africa.Ignoring these
population aspects might lead to artefacts andfalse-positive
results when searching for associations betweengenetic markers and
biomedical phenotypes, including diseaseassociation studies.
Furthermore, population genetic studies inNorth Africa should also
take this diversity into account. It isimportant to trace many
different variables when studying NorthAfrican populations, such as
the social and demographic char-acteristics of the studied groups
(if it is urban or rural, if it isgeographically isolated, etc.)
and their history (the Arab influencein the region, slave trade
routes, etc.). The genetic structure ofNorth African population is
determined by many different factorsapart from a simple ethnic
dichotomy Arab–Berber, and includingthose variables sheds lights in
the understanding of the popula-tions.
Related Articles
Reconstructing Human History Using Autosomal, Y-Chro-mosomal and
Mitochondrial MarkersGene Flow, Haplotype Patterns and Modern Human
OriginsGenetic Diversity in AfricaHumans: Demographic History
Glossary
Admixture The process of mixing of different populationswithin a
species.
Cultural diffusion The spread of cultural practices fromanother
group of people without genetic exchange.
eLS © 2017, John Wiley & Sons, Ltd. www.els.net 5
http://onlinelibrary.wiley.com/doi/10.1002/9780470015902.a0020819.pub2http://onlinelibrary.wiley.com/doi/10.1002/9780470015902.a0020795.pub2http://onlinelibrary.wiley.com/doi/10.1002/9780470015902.a0020800http://onlinelibrary.wiley.com/doi/10.1002/9780470015902.a0005077.pub2
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Genetic Heterogeneity between Berbers and Arabs
Demic diffusion The spread of a population in an
advancingwave.
Gene flow The movement of genetic variation from onepopulation
to another.
Haplogroup A group of similar allele combinations that share
acommon ancestor.
Inbreeding Reproduction that involves genetically
relatedindividuals.
Introgression The process of mixing of different species.
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Further Reading
Camps G (1974) Les civilisations préhistoriques de l’Afrique
duNord et du Sahara.
Hublin J-J and McPherron PS (eds) (2012) Modern Origins. A
NorthAfrican Perspective. New York: Springer.
Nielsen R, Akey JM, Jakobsson M, et al. (2017) Tracing the
peoplingof the world through genomics. Nature 541 (7637):
302–310.
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