of genetics: a contemporary
view*
Mesolithic
Neolithic LUP NUP EUP
U: 5.7%
HV: 5.4%
I: 1.7%
U4: 3.0%
H: 37.7%
H-16304: 3.9%
T: 2.2%
K: 4.6%
T2: 2.9%
J: 6.1%
T1: 2.2%
gets transmitted from generation to generation Hence, this new concept imposed the "cultural
alongside other mutations, which did occur ear- diffusion" model of Neolithisation in contrast to
lier. In other words, the mutation (a mistake in "demic diffusion" model advanced by Ammer-
the genetic text) became forever part of this text man and Cavalli-Sforza (1984).
and is transmitted together with all the mistakes The following decade witnessed a heated
that had appeared in this text earlier. When debate between two camps of geneticists, name-
comparing different texts (so-called haplotypes) ly the "cultural diffusionists" and the "demists".
it is possible to trace mutations back in time and Despite the ongoing debate, the methodological
reconstruct a "genealogy" of these texts. i.e. to limits and benefits of both models are apparent.
draw their "family tree". This tree is commonly The source database for demic diffusion model
rooted in the most recent common ancestor (a was much richer (hundreds of markers studied
"mitochondrial Eve") and each branch of the in dozens of populations) while Richards and col-
tree differs by its particular set of mutations. leagues were restricted to one marker (mtDNA)
Next, each twig of a certain branch carries all studied in a limited set of populations. Arguably,
mutations characteristic to this branch, togeth- as Barbujani and colleagues (1998) pointed out,
er with a set of additional "twig-specific" muta- the Palaeolithic origins of haplogroups found in
tions. These branches and twigs are called hap- Europeans do not necessarily imply that these
logroups (subhaplogroups) each of them unites haplogroups were present in Europe since the
a group of closely related haplotypes (which can Palaeolithic. As haplogroups age is calculated
be compared with a leaves of this tree). Assum- based on its diversity, they could have accumu-
ing an average rate of mutations one can calcu- lated diversity in other parts of the world ar-
late the age of each haplogroup by multiplying riving into Europe being already diverse. This
the number of accumulated mutations by the problem of the pre-existing diversity met ele-
mutation rate. gant solution in the following paper by Richards
This methodology, applied to the European and colleagues (2000), which became the most
mitochondrial pool (Richards et al., 1996), dem- recognized study of European genetics. In that
onstrated that most branches (haplogroups) paper the founder mtDNA lineages were identi-
found in Europe were much older that the Neo- fied which were deemed as the starting points
lithic and most of them fell into the age range of for the entire European diversity accumulated
the Upper Palaeolithic. Based on this evidence in situ. Although different criteria for "founding"
it was concluded, that European gene pool was resulted in slightly different time assessments,
formed by the initial peopling of the continent all calculations demonstrated the Upper Pa-
by anatomically modern humans (AMH) during laeolithic age for most European clusters of lin-
the Upper Palaeolithic, and that it is still pres- eages (haplogroups), while haplogroups whose
ent in the most of present-day Europeans. As appearance in Europe can be attributed to the
for the Neolithic expansion, it had, therefore, Neolithic period make up only a quarter of the
a limited impact on the European gene pool. total European gene pool (Fig. 2).
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frequent haplogroups. All but one of them came the database six times larger than previously
from the Near East: in the majority of cases possessed, comprising 20,000 European mtD-
during the Early Upper Palaeolithic (EUP) in NAs (Balanovska, Zaporozhchenko, Pshenich-
conjunction with the initial AMH dispersal and a nov, Balanovsky; MURKA Mitochondrial Data-
smaller part during the Neolithic epoch (in the base and Integrated Software, unpublished) we
course of Neolithisation, Richards et al., 2000). were able to recognise a much clearer patterning
The genetic landscape has been reshaped in the (Fig. 4). European populations altogether occu-
Mesolithic/Late Palaeolithic times, during the pying all parts of this plot provide a geometrical
repopulation of Europe from the southern Eu- illustration of the genetic variation in Europe.
ropean refugia. The only European haplogroup
that presumably had emerged in Europe (hap-
logroup V) became spread across the entire
continent during Mesolithic/Late Palaeolithic re-
colonisation (Torroni et al., 2001). The western
European origin of this haplogroup (the Franco-
Cantabrian refugium) is presumed, based on its
high frequency in this area as well as the oc-
currence of its phylogenetic predecessor (pre-V
lineages). However, a recent accumulation of
genetic data on previously poorly studied East-
ern Europe, enabled the present author (Bal-
anovsky, 2008) to suppose the occurrence of
an additional East European centre of origin of
this haplogroup. This finding is based on even
higher frequency and yet again, on the pres-
ence of pre-V lineages in East European steppe
area. Impossibility to distinguish between the
western and eastern European homelands em-
phasised the important feature of the European
mitochondrial landscape – its extreme homoge-
neity.
Indeed, when additional data from different
European populations became available the ge-
netic similarity in haplogroup frequencies (and
identity in haplogroup spectra) has been im-
mediately recognised (Simoni et al., 2000). As
a result, mtDNA studies has appeared dealing
with Europe as a whole, comparing it with the
Near East or other areas, while attempts to trace
genetic processes within Europe encountered
problems (Helgason et al., 2000). This devel-
opment was rather discouraging for archaeolo-
gists and linguists who were typically interested
in a genetic support of existing hypothesis in
their respective disciplines, although on a much
smaller scale. Fortunately, the paper entitled "In
search of geographic patterns in European mito-
chondrial DNA" (Richards et al., 2002) made the
point that with the emergence of a larger data-
Figure 4. Genetic relationships of European populations
set (with more than 3,000 individual mtDNAs) a from mitochondrial DNA perspective.
spatial structuring became more apparent (e.g. A. The multidimensional scaling plot (geometric distances
the south-north difference was acknowledged between points display the genetic distances between
among macro-regions of Europe: Mediterranean corresponding populations). Ellipses mark populations
belonging to the same linguistic group.
area, Central Europe, Scandinavia, and, surpris- B. The idealized approximation of the plot (A) by
ingly, the Basque Country). flower-like structure. Black core – proto-Indo-European
Presently one can affirm that size of the population; dotted ellipses – hypothetical extinct linguistic
dataset is the key factor. Having at our disposal groups.
9
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The most remarkable feature is that the pattern ru/main.html). This does not necessarily imply
distinctly resembles linguistic groups: popula- the Neolithisation (for example, major changes
tions cluster together according to their linguis- during the Bronze Age is one of alternative ex-
tic group of the Indo-European family. Three planations), but lends credence to the hypoth-
largest clusters are formed by Romanic, Slavic eses advocating a relatively recent origin (or at
and Germanic speakers, while Baltic and Celtic least late major reshaping) of the mitochondrial
speakers form smaller clusters and Albanians pool in Europe.
form a "cluster" of its own outside any other One may note that the significance of the
cluster. There are a few exceptions: Romanians, linguistic factor is quite obvious on the graph
Aromuns and Sicilians lie outside the Romanic (Fig. 4 A). However, the idea of a single proto-
cluster while Estonians join the Slavic cluster. In population totally depends on the flower-like
both cases the geographical distance (remote- structure of this graph (Fig. 4 B) and should be
ness for Romanians and Aromuns, proximity for therefore considered as one of the plausible hy-
Estonians) had probably a stronger impact on potheses.
genetics than the linguistic affiliation.
Among the non-Indo-European populations Y chromosomal landscape of the Europe
of Europe the Basques found their place outside
any other cluster but close to the Romance one While the "homogeneity" is the principal
(not surprisingly, considering the geographic feature of mitochondrial pool, the Y chromo-
proximity again). Finno-Ugric and Turkic speak- somal pool is characterized by a high heteroge-
ers are not shown on this plot because of their neity. As with mtDNA, there are seven Y chro-
extreme genetic variation, but on another plot mosomal haplogroups dominating in Europe.
they lie apart of Indo-Europeans. But while frequencies of mitochondrial haplo-
This linguistic structuring of European mi- groups are quite similar across Europe, Y chro-
tochondrial DNA follows a remarkable "flower mosomal haplogroups follow a clear geographi-
shape" pattern: all clusters looking like petals cal pattern (Fig. 5). Neither classical markers,
around the "core" of the flower. The possible ex- nor mitochondrial haplogroups demonstrated
planation of this pattern is that genetic and lin- such obvious and elegant trends. Therefore, Y
guistic differentiations were parallel processes chromosome became an effective instrument in
or, in better words, – two aspects of the same population genetics.
process, related to the multiplication and differ- One should remember that European gene
entiation of proto-Indo-European population in pool cannot be homogeneous and heteroge-
Europe. It is well known, that in many particular neous at the same time. The question is to what
cases distribution of genes is opposed to distri- degree different markers are able to reveal the
bution of language (especially in cases of the existing degree of variations. Having dozens of
language replacement by the elite dominance autosomal markers, classical population geneti-
model). However, in very general view, almost cists achieved reasonable resolution in assess-
all Europe is populated by speakers of one lin- ing the variation between populations (Cavalli-
guistic family and almost all Europe is geneti- Sforza et al., 1994). Mitochondrial DNA failed
cally homogenous. This allows speculations (like to reveal a difference between populations and
our flower-like interpretation of the genetic plot) successfully operates only at a higher hierarchi-
which consider genetic and linguistic evolution cal level: separating regions (Richards et al.,
as generally parallel processes, disregarding 2002) and linguistic groups (present study, fig.
partial exceptions. Such speculations inevitably 4). Y chromosome operates much better and
oversimplify both processes but could serve as a separates even subpopulations within the same
starting point for more detailed studies. ethnic group (Balanovsky et al., 2008). Recent
Therefore, one can accept as a working studies based on half of million autosomal mark-
hypothesis the differentiation of proto-Indo- ers became able to separate even individuals
European language into linguistic groups being within subpopulations (Novembre et al., 2008).
accompanied by genetic differentiation resulting This high differentiation power of Y chro-
in a clear clustering pattern (Fig. 4). This allows mosome (i.e. clear geographical clines of its
one to introduce time frames into the forma- haplogroups) was revealed already in the early
tion of the European mitochondrial landscape. It large-scale studies (Semino et al., 2000; Rosser
would coincide with origin and differentiation of et al., 2000). These clines have been recently
European branches of IE family, i.e. covers the summarized in a panel of frequency distribu-
last 5-6 millennia (Starostin et al., their linguis- tion maps (Balanovsky et al., 2008). Two main
tic database is avalable at http://starling.rinet. haplogroups, accounting altogether almost for a
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half of the total European Y chromosomal pool attributed to the Asian influence. Authors sup-
are distributed along the west-to-east axis. Hap- posed step-by-step migration from North China
logroup R1b accounts for roughly 50% of the Y to Eastern Europe, which started in early Ho-
chromosomal pool in Western Europe and de- locene and underwent a secondary expansion
creases eastward, while R1a reaches the same on its long way. Derenko and colleagues (2007)
high frequency in the east (Fig. 5) and decreas- studied microsatellite variation associated with
es westward. this haplogroup in more detail and tried to esti-
Analysis of another type of Y chromosomal mate its age. They identified two variants, one of
markers (microsatellite variation) also proved which migrated into Europe 6-10 ky ago, while
the western and eastern domains to be main the second (less frequent) variant was shown to
features of the Y chromosomal pool (Roewer et come by the way of a smaller and more recent
al., 2005). As it was stressed above, the "inter- migration, 2-4 ky ago. Although these time esti-
pretation by association" should be made with mations should be taken with great caution, the
caution. That is why attributing these domains both studies (Rootsi t al., 2007; Derenko et al.,
to Late Palaeolithic re-colonisation from two 2007) agree that north-east Europe had a sig-
principal refugia (the south-western and south- nificant (or even predominant) genetic legacy in
eastern ones) can be considered as a possible South Siberian/Central Asian populations.
but not yet proven hypothesis. (One of other This creates a problem for "two systems"
possibly hypotheses is attributing these genetic approach, because from mitochondrial perspec-
domains to descendants of Late Neolithic Bell tive Siberian/East Asian haplogroups appeared
Beaker and Corded Ware cultures). in low frequencies and only in the eastern edge
These two principal European haplogroups of Europe and did not account for a significant
R1b and R1a are shared between Europe and portion of the gene pool anywhere else in Eu-
other regions (Central Asia, Near East, India rope. (When low frequency of typical East Asian
and North Africa). But two other haplogroups, haplogroup F was found on Croatian isles and
I1 and I2a (according to previously used no- in even lower frequencies on Croatian mainland,
menclature the same haplogroups were labelled this was considered as a paradox and a spe-
as I1a and I1b, respectively) are restricted to cial paper (Tolk et al., 2001) tried to explain it
Europe, where they had likely originated. by possible medieval gene flow caused by trade
While R1b and R1a occupy the west and the routes of Venice). That is why a significant Asian
east, I1 and I2a predominate in the Europe’s presence in Europe, concluded from haplogroup
north and south, respectively. I1 which is fre- N1c remains one of the main inconsistencies be-
quent in Scandinavia and southern Baltic area tween Y chromosomal and mitochondrial genet-
has attracted less attention due to obviously late ic systems. From our point of view, this problem
colonization of this region. In contrast, the dis- could be resolved if one takes into consideration
tribution of haplogroup I2a (Balkan haplogroup) the fact that genetic boundary between Europe
has been widely debated. As southeast Euro- and Asia lies much eastern than Ural Mountains.
pean autochthonous haplogroup it could not be The western Central Asia (the Altai Mountains
attributed to Neolithic immigrants (or any other in particular) could be therefore considered as
immigrants) into Europe. We will discuss it in a genetically intermediary in present time and
more details below. primary "European" zone in the past. This view
Three remaining haplogroups (E, J, and explains why Y chromosomal haplogroup N (de-
N1c) are not evenly spread across the entire Eu- spite its origin in East Asia 20-30 ky ago) in pre
rope but are restricted to distinct areas. For this Neolithic or Neolithic times could be the char-
and other reasons they are believed to mark acteristic haplogroup for Caucasoid populations
later migration waves into Europe which did not in Eurasian steppe west from the Altai and also
cover the entire continent. for Mongoloid populations east from the Altai.
The haplogroup N1c (N3 or TAT, accord- From the western part of this area the haplo-
ing to previous nomenclatures) is restricted to group N1c could spread northward and north-
north-east Europe (mainly Finnic speakers) and westward by a number of migrations suggested
Siberia. During the last decade it remained un- for this area. This view also explains why these
clear whether is marks an eastward migration migrations did not bring East Eurasian mito-
from Europe or the opposite westward migra- chondrial haplogroups into Europe: the source
tion trend. In 2007 Rootsi and colleagues have area having mainly Western Eurasian haplo-
shown that this haplogroups could be deeply groups even in the contemporary gene pool. It
rooted in East Asian phylogeny and therefore the was more the case in earlier times before Turkic
occurrence of this haplogroup in Europe may be speakers brought East Eurasian haplogroups by
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their expansion into this area two millennia ago thors did not formulate this explicitly, their con-
onward. cept implies, that cultural diffusion took place
Two last Y chromosomal haplogroups to between regions (Analolia and Balkans, Balkans
be discussed are E and J. They predominate in and Central Mediterrania) while the demic diffu-
North Africa and Near East, and in Europe they sion occurred within regions.
are found mainly in Mediterranean area. Not all
sub-branches of these two haplogroups reached Ancient DNA
Europe, but mainly one branch of haplogroup
E (namely, E-V13) and two branches of haplo- Analysis of ancient DNA (aDNA) provides
group J (J-M241 and J-410). direct data on the former European gene pool,
While J-410 follows the separate pattern, which are free from assumptions and specula-
the other two haplogroups (and also haplogroup tions which often accompany deductions of past
I2a, mentioned above) are concentrated in the genetic processes, based on the contemporary
Balkans and have not been found in neighbour- genetic pattern. This advantage of aDNA may
ing regions with any significant frequencies. trigger a revolution in population genetics and
The ages of these haplogroups estimated from if this did not happen so far, this was due to
their STR diversity are: E-V13 from 4 to7,5 ky; limited quality and quantity of available aDNA
J-M241 from 3,5 to 6 ky; I2a from 5,5 and 10 ky evidence.
(Battaglia et al., 2008). This roughly coincides Problems with the quality (authenticity) of
with time of Neolithic transition in this part of aDNA data are dramatic because of the possible
Europe. The model suggested by Battaglia and contamination by modern DNA. For this reason
colleagues states that Neolithic cultural package some aDNA results may be false, and many
was adopted by local Mesolithic populations of early aDNA papers were criticized exactly from
the Balkans which started grow in numbers, ex- this point of view. This problem could be par-
panding farming across the entire Balkan penin- tially resolved in few high-standard laboratories
sula and later transmitting this package to other only, which have special equipment to minimize
Mesolithic populations of Europe. This model ex- risk of contamination. Cross-checking, i.e. in-
plains why these three haplogroups are restrict- dependent analysis of the same ancient sample
ed to the Balkans, why they exhibit decreasing in different aDNA labs is the second condition.
frequency towards other part of Europe and why The third one is implementing the "modern DNA
their age is similar to that of the Neolithic transi- free" style of excavation into the practice of the
tion. archaeological fieldwork. Having these three
However, the internal logic of this model is conditions met, one can reach reasonably de-
opposite to those applied by Richards and col- gree of authenticity of the aDNA results.
leagues in relation to mitochondrial DNA. In- The quantity problem consists in the scar-
deed, Richards and colleagues proved that mi- city and limited sample sizes of the aDNA data.
tochondrial haplogroups whose diversity was Again, this problem could be solved only par-
accumulated in Europe in situ are of Palaeoli- tially, by increasing the number of aDNA studies
thic age; and from this fact they concluded that and average sample size per study. Fortunately,
present-day Europeans are descendants of Pa- both factors tended to increase in the last de-
laeolithic population of Europe (Richards et al., cade, still trace amounts and high fragmenta-
2000). Eight years later, Battaglia and colleagues tion of ancient DNA samples hinder its high-
proved that Y chromosomal haplogroups whose throughput analysis.
diversity was also accumulated in Europe in situ Because of these limitations aDNA at least
are of Neolithic age; but from this contrasting presently cannot be the main source of genetic
fact they concluded also that present day Euro- knowledge about the Neolithisation. But it is
peans are descendants of Palaeolithic population already one of the important sources on this
of Europe (Battaglia et al., 2008). Both studies problem. Indeed, analyses of Neandertal mito-
are reasonably substantiated and their conclu- chondrial DNA (Krings et al., 1997; Ovchinnikov
sions look correct. However this example illus- et al., 2000), though being criticized for prob-
trates that genetic studies need more robust and able mistakes in sequencing, put an end to a
universal logic, at least when dealing with such lengthy discussion of the possible assimilation
complex process like the Neolithisation. In this of the Neandertal populations by anatomically
particular case the possible logical compromise modern humans. Specificity of Neandertal mi-
lays in the fact that concept of Battaglia and co- tochondrial type (Currat, Excoffier, 2004) and
authors actually implies both, cultural diffusion absence of this type in present day Europeans
and demic diffusion models. Although the au- (Behar et al., 2007) allow to root European gene
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pool in AMH colonization of the Europe, disre- and quality of aDNA data which might allow bet-
garding the previous epochs. ter chronological and geographical resolution of
Direct genetic data on first Neolithic groups genetic processes in the near future.
in Europe are of course most promising source
for choosing between the demic and cultural dif- Conclusions
fusion models of Neolithisation. Such data are
now available for Neolithic population of the The increasingly accumulating genetic data
Iberian peninsula (Sampietro et al., 2007) and on extant and extinct (aDNA) European popu-
Neolithic population of the Central Europe (sites lations are most frequently discussed in terms
of Linear Band Ceramic with age of 7.0 – 7.5 of two opposite concepts: demic diffusion and
ky; Haak et al., 2005). Iberian Neolithic popula- cultural diffusion models of Neolithisation. In
tion was shown to be genetically similar to the hands of Cavalli-Sforza and his colleagues the
present day Iberian population. In contrast, LBK genetic mirror reflected Neolithic expansion
population in Central Europe was shown to be across Europe (demic diffusion); but in hands of
genetically distinct from the present day popu- present-day writers this mirror reflects mainly
lation of that or any other region of Europe). the Palaeolithic legacy of Europeans and cultural
The most remarkable feature of the Neolithic diffusion model is needed to explain spread of
population was mitochondrial haplogroup N1a farming.
found in 6 of 24 individuals. This haplogroup Understanding the genetic history of Eu-
is virtually absent in present-day Europe. The rope implies clarifying relative significance and
mathematical simulation has shown that if this patterns of each of the following processes:
Neolithic population was source of present-day 1. the initial dispersal of AMH in Europe (Upper
Europeans they could not have lost this haplo- Palaeolithic);
groups by stochastic genetic drift. 2. the restructuring of the genetic landscape
It was therefore concluded (Haak at el, during the Mesolithic repopulation of the
2005) that Neolithic LBK population did not be- Europe from two-four refugia;
come parental for present-day European gene 3. the importance of the Neolithic expansion
pool, but became dissolved in pre-existing Eu- viewed as the spread of early farming com-
ropean populations. This conclusion is therefore munities or spread of Neolithic cultural
in agreement with cultural diffusion model in as- package;
suming that since Neolithic farmers arrived in 4. the role of post-Neolithic human movements
Europe, the farming was adopted by aboriginal within Europe;
populations and first farmers did not leave any 5. the "oriental" influence in different epochs –
considerable genetic legacy in their new home- from Palaeolithic to Medieval times.
land. To address these questions population ge-
The study by Haak and colleagues did an- netics operated with autosomal (classical) mark-
swer the question: "what happened with first ers in the past and autosomal (DNA) markers
farmers after their arrival in Europe". To address may became the new standard in the future,
the another question, "where these first farm- while the present day studies are based on mi-
ers came from", the consequent study was per- tochondrial DNA and Y chromosomal variation.
formed (Haak, pers. comm.). Based on extend- Analysis of mtDNA demonstrated that most
ed dataset (44 individual mtDNAs from different of European haplogroups came from the Near
sites of early LBK culture) it was found that this East during the Upper Palaeolithic times and
population is genetically similar to present day Neolithic migration of Near Eastern farmers did
populations of Northern Mesopotamia, southern not contribute much into the European gene
Caucasus and eastern Anatolia. Although the pool. The south-east – northwest cline within
genetic composition of this area could be dis- Europe, as established by many genetic mark-
turbed after the Neolithic period by subsequent ers, is not considered anymore as the trace of
migrations, it is reasonable to suppose that in- Neolithic expansion, because Palaeolithic colo-
ner areas of the Near East were homeland for nists used likely the same geographical route.
migrating groups who finally brought these mi- Y chromosomal data reveal distinct do-
tochondrial lineages into the LBK population of mains of prehistoric movements within Europe.
the Central Europe. Particularly, two different haplogroups predomi-
Of course, this data give rise to many new nate in Western versus Eastern Europe, and one
questions, and currently available aDNA data may speculate about two secondary homelands,
are not sufficient to address them. The mod- associating them with Mesolithic refugia or cen-
erate optimism is based on increasing number tres of later expansions.
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Southeast Europe (the Balkans) which de- D.Primorac, R.Hadziselimovic, S.Vidovic, K.Drobnic,
serves special attention as gates into Europe, N.Durmishi, A.Torroni, A.S. Santachiara-Benerecetti,
P.A. Underhill and O. Semino (2008) "Y-chromosom-
is populated by three different Y chromosomal al evidence of the cultural diffusion of agriculture in
haplogroups exhibiting similar patterns: being southeast Europe", European Journal of Human Genet-
autochthonous for Europe these haplogroups ics Advance online publication 24 December 2008; doi:
started to expand in time frames comparable 10.1038/ejhg.2008.249
Behar D.M, S. Rosset, J. Blue-Smith, O. Balanovsky, S.Tzur,
with the Neolithisation; it was supposed that D. Comas, R.J. Mitchell, L. Quintana-Murci, C.Tyler-
this expansion might took the form of Balkan’s Smith, R.S. Wells and Genographic Consortium. (2007)
Mesolithic population adopting farming from "The Genographic Project public participation mitochon-
their Anatolian neighbours. drial DNA database" PLoS Genetics Vol 3(6):e104.
Cavalli-Sforza L.L., P. Menozzi and A. Piazza A. (1994) His-
Analysis of ancient DNA indicated that first tory and Geography of Human Genes. Princeton: Princ-
Central European farmers (LBK) were of Near eton University Press. 1059 p.
Eastern origin but did not left recognisable de- Chikhi L, R.A. Nichols, G. Barbujani and M.A. Beaumont
scendants. The early farmers in Iberia (and pos- (2002) "Y genetic data support the Neolithic demic dif-
fusion model", Proceedings of the National Academy of
sible in other areas of late Neolithisation) were Sciences USA Vol 99(17):11008-13.
of aboriginal European genetic type. Childe V.G. 1928. The Dawn of European Civilization. New
Genetic mirror shows a controversial pic- York, Knopf
ture: even in this summary "indigenous" Balkan Currat M, and L. Excoffier (2004) "Modern humans did not
admix with Neanderthals during their range expansion
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cally the European gene pool and consequently Haak W, P. Forster, B. Bramanti, S. Matsumura, G. Brandt,
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