Where Did The European Men Come From

Kalevi Wiik
on the development of the human family, specifically the

European part of the family, over the last 40,000 years.
.
Nearly all European men can be classified into one of the

nine most frequently occurring Y-chromosome haplo- The genetic development and migrations of the ancestors
groups or clans. Haplogroups are defined by mutations of European men can be presented in the following ten
or single nucleotide polymorphisms (SNPs). The nine phases; see .
most common haplogroups, in alphabetical order, are
E3b, G, I1a, I1b1-P37, I1b2-M223, J2, N3, R1a, and (1) About 50 thousand years ago (kya) all the ancestors
R1b. Sometimes, two more haplogroups, E3a and (in of the present European men still lived in northeastern
Northeast Europe) N2, are added to the list, which Africa and formed only one clan.
makes the total number eleven. In addition, there are
many other rare haplogroups in Europe. (2) About 45 kya the clan was split into an “African”
Clan E and an “Asian” Clan F, and the “Asian” clan
shows the Y phylogenetic tree for European moved out of Africa to the Arabic peninsula and the
haplogroups. It includes forty haplogroups attested in Near East.
Europe, but thirty of these are so rare that their frequen-
cies do not reach one per cent anywhere in Europe. The (3) About 40 kya part of the “Middle Eastern” Clan F
six thicker branches or groups of branches of the tree, gave rise to the “Central Asian” Clan K.
representing Haplogroups R1, I, N, E, J, G, and some of
their subgroups, are discussed in this review. The more (4) About 35 kya two new clans, R and NO, branched
detailed structure of these haplogroups may be seen in off from K. Clan R moved to western Central Asia and
the Y phylogenetic tree maintained by ISOGG (2007). Clan NO to eastern Central Asia.
The tree in is a simplification of that in the
article by Underhill and Kivisild (2007). (5) About 30 kya Clan R was split into R1 and R2, and
Clan R1 moved to the steppe area between the Ural
The SNP’s presently available provide information only mountains and the Caspian Sea.
on the large-scale classification of Y chromosomes. Each
haplogroup may be further resolved into clusters based (6) About 25 kya one branch of Clan R1, Clan R1b,
on similarity of Y Short Tandem Repeats or Y-STR’s. reached Iberia and the Atlantic Coast, and somewhat
Additional information may be derived from other ge- later Clan R1a branched from R1 and became common
netic markers. However, only the SNP classifications in the present-day Ukraine.
will be used in this paper, but this provides information
____________________________________________________________
(7) About 25 kya the “Middle Eastern” Clan F sent
another branch to Anatolia and further to the Balkans,
Address for correspondence: Kalevi Wiik, kalevi.wiik@pp1.inet.fi
and a new sub-Clan I emerged.
Received: 10 Oct 2007; accepted: 03 Feb 2008.
35
36
A phylogenetic tree of about forty Y-chromosome haplogroups and subhaplogroups in

Europe and its vicinity. The tree is a simpificaton of the Y-chromosome haplogroups of the entire
world. The subhaplogroups of I have been regrouped. Source: Underhill-Kivisild (2007).
The migration routes of the ancestors of European men from Africa to Europe. Grey routes show the
genetic developments (by mutations) of European Haplogroups E3b, F, G, I, j, N, and R1. Coloured circles =
the approximate geographic locations of the main four male populations during the last glacial maximum (LGM)
about 20 kya. Red, blue, brown, and yellow arrows show essential population movements during the
recolonization of northern europe during the late glacial period (about 16-10 kya). Green arrows indicate arrival
of Early Farmers in the Balkans and Mediterranean coast about 10 kya.
Wiik: Where Did European Men Come From? 37
(8) Perhaps at about the same time, Clan NO in eastern “Old Europeans,” rather the Early Farmers. The rather
Central Asia developed into “Siberian” Clan N and rare Clan F* in Europe may belong to either group. If
moved towards the north. we think the men of F* clan left the Middle East for the
Balkans before the emergence of effective farming, they
(9) Clan N was split into two sub-clans, N3 and N2, and are comparable to the men of Clan I and belong to the
these moved first to northwestern Siberia and later to Old Europeans, but if they left the Middle East only after
Eastern and Northeastern Europe. the beginning of farming, they are comparable to the
men of Clans E3b, J and G and represent the Early
After these nine phases, Europe experienced a cooling Farmers.
climate and the onset of the Last Glacial Maximum
(LGM). During the LGM the ancestors of the European
men retreated from northern Europe into four refuges
located in Iberia, the Ukraine, the Balkans, and Siberia. European men (more precisely, their Y chromosomes)
These core areas were habitable even during the coldest can be classified into two categories:
periods of the LGM.
(a) Those who are Old Europeans in the sense that, at
(10) About 10 kya the farmers of the Middle East, the start of the LGM, their paternal lineages already
representing African Clan E (its sub-clan E3b) and two were in Europe and they came to the four refuges when
sub-clans of F (the “Caucasian” Clan G and the “Near they were forced out of northern Europe. They were
Eastern” Clan J), spread to Anatolia and further to first to repopulate Europe after the LGM and they
Greece and the Mediterranean coast. formed the bulk of the present European male popula-
tion.
Clans I, E3b, J, and G all originate from the Middle East,
but only E3b, J, and G (not I) belong to the group of (b) Those who during the Ice Age still were in the warm
“Early Farmers.” Clan I had spread into Europe before regions of Asia and Africa. These latter populations
the emergence of effective domestication of wild plants came to Europe during the Neolithization of Europe (the
and animals (i.e. the beginning of agriculture and cattle arrival of farming) that started about 10 kya. The fre-
raising) in the Middle East. Because of their early quencies of the two groups of men are shown in the
departure, they were still hunter-gatherers at the time of tables in columns OE (Old Europeans) and EF (Early
the arrival of the Early Farmers in the Balkans, and they Farmers). The frequency of the Old Europeans will be
were taught to cultivate land and raise cattle by their considered here as the sum of R1b + R1a + I + N and
”Middle East brothers” after the ”reunion of the fami- that of the Early Farmers is the sum of E3b + J2 + G.
ly” in the Balkans. Accordingly, Clan I represents the
Four refuges and their typical Y-chromosome haplogroups. From west to east they are the
Iberian, Balkan, Ukrainian, and Siberian refuges. The bigger circles represent the four refuges, and
the smaller ones the peak areas of the corresponding populations today.
38
smaller circles represent locations of the peak areas of

the four populations today (Clan I has two peak areas in
The European populations are generally thought to have the map). Each refuge had its own prominent Y-chro-
been forced southward into three refuges during the Last mosome haplogroup: the haplogroup was R1b in Iberia,
Glacial Maximum (about 20 kya and thereafter). The I in the Balkans, R1a in the Ukraine, and N3 in Siberia.
refuges were (from the west to the east) in Iberia, the Until about 10 kya, the European male population
Balkans, and the Ukraine. In the present study, one consisted typically of only these four clans: R1b, I, R1a,
additional refuge, that of Siberia, is considered. The and N3. There may have been others, but their impact
four refuges are seen in as the larger circles; the on the European male population was smaller.
Population Old Europeans Early Farmers Other

R1b R1a I N3 Total E J G Total Total
1 Andalusian 65.5 3.4 68.9 10.3 13.3 23.6 7.5
2 Basque-Sp 88.9 6.6 95.5 2.2 2.2 4.4 0
3 Basque-Fr 86.4 9.1 95.5 4.5 4.5 0
4 Catalan 79.2 4.2 83.4 4.2 4.2 8.3 16.7 0
5 French 52.2 17.4 69.6 8.7 17.3 26.0 4.4
6 Dutch 70.4 3.7 22.2 95.9 3.7 3.7 0.4
7 German 50.2 6.2 37.5 93.7 6.2 6.2 0
8 Czech + Slov 35.6 26.7 15.6 2.2 77.9 2.2 8.9 4.4 15.5 6.6
9 Italian 62.0 4.0 8.0 70.4 2.2 14.0 10.0 26.2 3.4
10 Calabrian 32.4 32.4 16.4 32.4 8.0 56.6 11.0
11 Sardinian 22.1 37.7 59.8 10.4 10.4 14.2 35.0 5.2
12 Croatian 10.3 29.3 44.8 84.4 6.9 5.2 1.7 13.8 1.8
13 Albanian 17.6 9.8 19.6 47.0 23.6 27.5 2.0 53.1 0
14 Greek 27.6 11.8 7.9 47.3 23.7 22.3 2.6 48.6 4.1
15 Macedonian 10.0 35.0 20.0 65.0 15.2 20.0 35.0 0
16 Polish 16.4 56.4 23.6 96.4 3.6 3.6 0
17 Hungarian 13.3 60.0 11.1 74.4 8.9 2.2 2.2 13.3 2.3
18 Ukrainian 2.0 54.0 18.0 6.0 74.0 4.0 6.0 4.0 14.0 12.0
19 Georgian 14.3 7.9 22.2 36.5 30.1 66.6 11.2
20 Turkish 6.6 6.6 3.3 3.3 19.8 13.3 43.3 6.6 63.2 17.0
21 Lebanese 6.4 9.4 3.2 19.3 25.8 35.1 3.2 64.1 16.6
22 Syrian 15.0 10.0 5.0 30.0 10.0 45.0 55.0 15.0
23 Saami 8.3 8.3 41.7 41.7 100 0 0
24 Udmurt 11.6 37.2 7.0 30.2 86.0 4.7 4.7 5.4 8.6
25 Mari 13.0 4.3 69.5 86.8 6.5 6.5 6.7
agriculture expanded as the indigenous people acquired

a new knowledge of how to cultivate land and they were
The situation changed about 10 kya when agriculture not moving away from their old living areas. According
started to expand from the Middle East. The new to the demic diffusion model, this new form of subsis-
subsistence system expanded into Europe in two differ- tence was brought to Europe mainly by three clans: E3b,
ent ways: (a) as demic diffusion, according to which the J2, and G2. shows the relative portions of these
new subsistence system was brought to new areas by clans in southwestern Asia (e.g. the Middle East) and
newcomers who moved in and brought agriculture with Europe. The percentages of the four haplogroups of the
them, and (b) by cultural diffusion, according to which Early Farmers (E3b, J2, JxJ2, and G2) in relation to the
The Early Farmers of four haplogroups: E3b, J2, J x J2 (J, not in J2--probably all J1), and G2
in Europe. The white section of each circle belongs mainly to the Old European populations R1b, I, R1a,
and N3, while the sum of the coloured sections show the approximate share of the Early Farmers. The map
is based on King and Underhill (2002).
percentages of the haplogroups of the populations from important sources for Clans E3b, J2, and JxJ2 (probably
the four refuges (R1b, I, R1a, and N3) can, at least to J1). J2 came in about equal portions from the Caucasus
some extent, be used as an indicator of how large the and Lebanon, with slightly less from Syria.
early farming populations arriving in various parts of
Europe were. I call the two groups of haplogroups and
clans “Early Farmers” and “Old Europeans.” The
percentages of the two groups can be seen in and In many cases the Wikipedia and Balanovsky maps
. contain enough data to indicate where the European
men have come from. In some cases, wider maps show-
The table and maps can be summarized as follows: ing the whole of Eurasia are needed for this purpose.
Next, I consider four groups from a wider perspective:
(1) In the Middle East and Anatolia, the Early Farmers how far do the distributions of certain haplogroups
represent the majority (about 55-64%), while in Europe extend outside Europe?
the frequency declines sharply in a clinal gradient from
about 35-57% of the male populations in the southern (1) Haplogroup R1b has its peak values in West Europe
Balkans and southern Italy, to a frequency near zero in and its total area extends far beyond the eastern border
far northwest Europe. of Europe. In the distribution of R1b is seen to
extend as far east as Uiguria (in northwestern China).
(2) The coloured slices of the easternmost three circles in The fact that this haplogroup has two secondary peaks
show that the Caucasus was a major source area outside Europe (one in Georgia and the other in Uiguria)
for Clans J2 and G2 and that Lebanon and Syria were
40
The geographic distribution of Haplogroup R1b. Source:

Wikipedia.
tends to indicate that the R1b men may have arrived in subgroup stayed in the Ukrainian refuge during the cold
Europe from the Caucasus or Central Asia. periods of the Ice Age. The Eurasian total area of
Haplogroup R1a is seen in . The haplogroup has
(2) Clan R1a, the “brother clan” of R1b, has a very wide four peak areas: one in northern India, another in Altai,
distribution with four peak areas. The East-European a third one in the Mari area (Central Volga region), and
The total geographic distribution of Haplogroup R1a. Source:

Kivisild (2005).
The main areas of concentration of The eastern extentions of Haplogroup R1a

Haplogroup R1a. The map also shows the in India and Mongolia. The map is based on a map
development of Haplogroup R1a from Haplogoup in Sahoo et al. (2006).
F (through intermediate stages K and P).
a fourth one in the Polish-Russian region; see also (4) The high frequencies of Haplogroup N in northeast-
. The two European maximum areas are also seen in ern Europe and practically all Siberia show that the men
detail in later in this article. of this clan came to Europe from the Siberian refuge; see
. This group can also be called “Uralic-Altaic” as
(3) Haplogroup I is restricted for the most part to many of its present-day representatives speak Uralic
Europe. An indication of the direction of arrival of this (Finno-Ugric) or Altaic languages. Group N was divided
clan is seen in in which the frequencies are into two main parts, N3 and N2; see
relatively high (4-12%) in Anatolia. . Clan N2 was to inhabit northernmost Siberia, while
The geographic distribution of all the subhaplogroups of N.

Source: Rootsi et al. (2006).
Frequencies of Haplogroup N3 in Eurasia.

42
Frequencies of Haplogroup N2 in Euraasia.
The development of Haplogroup N from F through K and

NO and the potential routes of the clans in question.
The geographical distribution of Haplogroup E (including all of its subhaplogroups).
Clan N3 extended to the west and east; today, Clan N2

speaks Samoyedic languages, while the western clans of
N3 speak Finno-Ugric languages and eastern clans
Altaic and Paleo-Siberian languages.
(5) From the European perspective, Haplogroups E3b,

J2, and G belonged to the first farmers to arrive in
Europe from the Middle East.
(a) Haplogroup E3b is a branch of the ”African” Haplo-

group E; see .
(b) The source area of the ”Near Eastern” Haplogroup

J was perhaps on the eastern coast of the Mediterranen;
its subhaplogroup J2 has its maximum area today in the
Caucasus and Anatolia; see .
(c) The source area of Haplogroup G was perhaps in

Pakistan and India; from the European point of view, its
peak values are today in the Caucasus; see .
Haplogroup E originated in Africa. This is true also of

its European subhaplogroup E3b. In Africa, this sub-
group has four separate peak areas; these are in South-
ern Africa, Morocco, Northern Libya, and the Horn of
Africa. The last two bear witness to the route from Frequency zones of the ”European”
northwest Africa through Anatolia to Europe. (The subhaplogroup E3b in parts of Africa, Asia, and
frequencies are relatively high also in Anatolia.) In Eu-
Europe. A southern peak area of E3b in southern
rope, Haplogroup E3b has peak values (about 25%) in
Albania; about equally high frequencies are also typical
Africa is not visible on the map.
44
The geographic distribution of Haplogroup J2. Source: Sengupta et al. (2006).
of some other parts of the Balkans and southern Italy.

Many of the newcomers representing subclan E3b learnt
agriculture on their way through the southwestern part
of the Middle East. They represent one group of the
typical Early Farmers of Europe.
It is likely that there is at least some interdependence

between the genome and languages in Europe. The
languages spoken in Europe belong to four independent
and unrelated phyla: Indo-European, Basque, Finno-Ug-
ric (Uralic), and Turkic (Altaic). The distribution of
these phyla is shown in
The Indo-European languages represent seven groups:

Germanic, Baltic, Slavic, Romance, Celtic, Albanian,
and Greek. The Finno-Ugric languages belong to five
groups: Finnic, Volgaic, Permic, Ugric, and Samoyedic
(represented in Europe only by Nenets). The Turkic Four language phyla in Europe. Blue =
languages in Europe are Tatar, Chuvash, Bashkir in the Indo-European, yellow = Finno-Ugrian (Uralic),
Volga-Ural area, and Turkish in Turkey. The distribu- grey = Turkic (Altaic), and red = Basque.
tion of the language groups or families is shown in
.
The question of which language a population speaks

today may be more dependent on the languages spoken their mates. Accordingly, the languages of men have
in the past by ancient men than by those spoken by usually remained unchanged while women have some-
ancient women. This statement is supported by the idea times accepted a new language in their new living area.
that women were more mobile than men—the men have If this is true, Y-chromosome data being reviewed here
supposedly more often stayed in their original living may provide more information about the spread of
areas, while women have often moved to the homes of ancient languages than mitochondrial DNA.
The approximate areas of European language groups:

(1) The Indo-European languages: Blue = Germanic (CG = Continental Geramnic, BG = Britain Germanic, and
NG = North Germanic), B = Baltic, dark green = Slavic (WS = West Slavic, ES = East Slavic, and SS = South Slavic),
C = Celtic, R = Romance, A = Albanian, G = Greek, O = Ossetian.
(2) The Finno-Ugric (Uralic) languages: F = Finnic, Sa = Saami, V = Volgaic, P = Permic, H = Hungarian (a Ugric
language), N = Nenets (a Samoyed language).
(3) The Turkic languages: Ta = Tatar, Ch = Chuvash, Bsh = Bashkirian, Tu = Turkish.
(4) The Basque language: Bs.
model used than by genetic reality. The maps represent-

ing approximate copies of those in Balanovsky et al. are
different in that they are more reliably based on real
The maps that follow show the approximate frequencies genetic data. The maps are shown in pairs so that the
of ten main Y-chromosome haplogroups in various parts map on the left is a Wikipedia map and that on the right
of Europe. In some cases haplogroups of more restricted a map from Balanovsky (2008). None of the maps are
distribution are also considered. The maps are based on exact copies from the original sources but have been to
the synthetic maps of Wikipedia (in some cases im- some extent reshaped by the present author. To help the
proved by the results of some recent well known genetic reader place each haplogroup in its proper ice age ref-
studies), and on those in Balanovsky et al. (2008). The uge, the main Haplogroups R1b, R1a, I, and N will be
next few maps are presented in pairs so that each pair referred to respectively, as “Iberian,” “Ukrainian,"
represents one haplogroup with the map on the left “Balkan," and “Siberian” haplogroups.
based on the Wikipedia version and that on the right on
Balanovsky et al. The copyright of the original Wikipe-
dia maps is owned by Relative Genetics, Inc.
From this point Europe is treated as eight separate
regions: (1) Central Europe, (2) North Europe, (3) East
Europe, (4) the Balkans, (5) the Italian Peninsula, (6)
Unfortunately, the two groups of maps (Wikipedia and Iberia, (7) the Atlantic Coast of Europe, and (8) the
Balanovsky) are not quite analogous in detail. In the British Isles; see . The male populations of each
Wikipedia maps, the isoglosses and zones of haplogroup region are mapped from the point of view of the follow-
frequencies are more regular but less reliable; in some ing haplogroups: (1) R1b, (2) R1a, (3) I, (4) N, and (5)
cases, they seem to be caused more by the mathematical E3b+J2+G.
46
Frequencies of the Iberian Haplogroup R1b in Europe.
Frequencies of the Ukrainian Haplogroup R1a.
Frequencies of the Scandinavian Haplogroup I1a, a aubhaplogroup of the

Balkan Haplogroup I.
Frequencies of the German Haplogroup I1b2-M223,

subhaplogroup of the Balkan Haplogroup I.
Frequencies of the West Balkan Haplogroup I1b1-P37, subhaplogroup of

the Balkan Haplogroup I.
Frequencies of the Northeast European Haplogroup N3, subhaplo-

group of the Siberian Haplogroup N. The N3 area extends to Siberia and reaches its
peak values (about 85%) in the Yakuts near the Pacific Ocean.
48
Frequencies of the Samoyed Haplogroup N2,

subhaplogroup of the Siberian Haplogroup N.
Frequencies of the Early Farmers Haplogroup E3b, subhaplogroup of

the African Haplogroup E.
Frequencies of the Early Farmers Haplogroup J2, subhaplogroup of the Near Eastern
Haplogroup J.
Frequencies of the Early Farmers Haplogroup G in Europe.
The total area of Central Europe is shown in

which also shows the areas of the seven Central Euro-
Linguistically, Central Europe consists of a Germanic, pean states, as well as the approximate locations of the
Slavic, and Finno-Ugric area. The Germanic language is populations involved in this study. Eleven populations
German (spoken in Germany, Austria and Scwitzer- are considered in Germany, five in the Czech Republic,
land); the West Slavic languages are Polish, Czech, and eight in Poland, three in Austria, and two in Switzerland.
Slovak; the Finno-Ugric language is Hungarian. Slove- The other Central-European states (Slovakia and Hun-
nian is sometimes (not, however, in this study) included gary) are represented only by one overall national popu-
in Central Europe; it then represents a fourth language lation each, while Austria and Switzerland are
group, the South Slavic languages. represented by both an overall national population and
two or three regional populations. The haplogroup
frequencies of the Germans and Poles are from Kayser et
al. (2005), and those of the Czechs from Luca et al.
(2007).
is a compilation of haplogroup frequencies for

the different populations. The Polish, German, and
Czech parts of the table are simplifications of the equiv-
alent tables in Kayser et al. (2005) and Luca et al.
(2007). The frequencies of the three Austrian popula-
tions and two Swiss populations were calculated by Kari
Hauhio using YHRD-data processed through Whit
Athey’s haplogroup predictor (Athey, 2005; Athey,
2006). The subhaplogroup frequencies from the pro-
gram were added to correspond to the major haplo-
groups shown in .
bear witness to the fact that agriculture

arrived in Central Europe from two directions, the Bal-
The eight European subareas dealt with kans and Western Europe. Agriculture had first arrived
from the Middle East and Anatolia in Greece and it
separately in this article. CE = Central Europe, NE
continued from there as two branches: one came to
= North Europe, EE = East Europe, B = the Central Europe by land, while the other extended from
Balkans, It = the Italian peninsula, Ib = Iberia, WC Greece along the Mediterranean coast and the Atlantic
= the Western Atlantic area, and Br = the British coast of West Europe by boat.
Isles.
50
R1b R1a1 I N3 OE E3b DE J2 G EF F P K Other n
1 Wroclaw 12.9 48.5 12.9 5.0 79.3 11.9 0 2.0 13.9 5.9 1.0 0 0 101
2 Warsaw 17.4 54.5 19.0 1.7 92.6 2.5 0.8 3.3 6.6 0.8 0 0 0 121
3 Lublin 12.5 62.5 11.6 0.9 87.5 3.6 0.9 3.6 8.1 2.7 0 1.8 0 112
4 Gdansk 7.3 60.0 21.3 3.3 91.9 3.3 0 2.7 6.0 1.3 0 0.7 0 150
5 Krakow 8.0 64.0 15.0 4.0 91.0 3.0 0 2.0 5.0 2.0 2.0 0 0 100
6 Szczecin 11.4 53.3 21.9 3.8 90.4 6.7 0 1.9 8.6 1.0 0 0 0 105
7 Suwalki 7.3 56.1 15.9 11.0 90.3 2.4 0 2.4 4.8 3.7 0 1.2 0 82
8 Bydgoszcz 14.8 55.6 18.3 2.8 91.5 3.5 2.1 2.1 7.7 0 0 0.7 0 142
Polish total 11.6 57.0 17.3 3.7 89.6 4.5 0.5 2.5 7.5 2.0 0.3 0.5 0 913
1 Berlin 23.3 22.3 32.0 1.9 79.5 9.7 0 1.9 11.6 3.9 0 3.9 1.0 103
2 Leizig 43.1 27.1 14.6 0.7 85.5 6.9 0 2.8 9.7 3.5 1.4 0 0 144
3 Magdeburg 34.0 21.0 25.0 1.0 81.0 7.0 0 2.0 9.0 6.0 3.0 1.0 0 100
4 Rostock 32.3 31.3 22.9 2.1 88.6 6.3 0 2.1 8.4 2.1 0 1.0 0 96
5 Greifswald 37.5 19.2 24.0 1.0 81.7 2.9 0 2.9 5.8 3.8 5.8 1.9 1.0 104
6 Hamburg 37.9 16.8 31.7 1.9 88.3 0 0.6 5.0 5.6 3.7 1.2 1.2 0 161
7 Muenster 37.3 7.8 26.5 1.0 72,6 9.8 0 4.9 14.7 7.8 1.0 2.9 1.0 102
8 Freiburg 54.9 10.8 16.7 0 82.4 4.9 0 8.8 13.7 2.9 1.0 0 0 102
9 Cologne 41.7 15.6 19.8 6.3 83,4 5.2 0 5.2 10.4 2.1 1.0 3.1 0 96
10 Mainz 44.2 8.4 22.1 1.1 75.8 11.6 2.1 6.3 20.0 3.2 0 0 1.1 95
11 Munich 41.1 14.3 23.2 0.9 79.5 7.1 0 2.7 9.8 8.0 0 2.7 0 112
Ger. total 38.9 17.9 23.6 1.6 82.0 6.2 0.2 4.0 10.4 4.3 1.3 1.6 0.3 1215
1 Klatovy 22.9 35.4 25.1 0 83.4 4.2 2.1 8.4 14.7 0 2.1 48
2 Pisek 29.2 29.2 24.6 3.1 86.1 1.5 4.5 6.2 12.2 1.5 0 65
3 J.Hradec 26.5 32.7 14.3 2.0 75.5 8.2 2.0 6.1 16.3 0 8.1 49
4 Trebic 32.7 34.7 10.2 2.0 79.6 6.1 8.1 4.1 18.3 0 2.0 49
5 Brno 28.3 41.3 15.2 0 84.8 6.5 6.5 0 13.0 0 2.2 46
Czech total 28.0 34.2 17.9 1.6 81.7 5.1 4.7 5.1 14.9 0.4 2.9 257
22 40 17 3 82 10 3 0 13 0 2 0
31.8 14.0 38.8 1.6 86.2 13.2 0 13.2 0 0 0.8
20 26 26 72.0 11 8 2 21.0 2
1 Tyrol 59.4 12.5 6.2 78.1 15.6 6.3 21.9

2 Graz 14.3 42.9 28.6 85.8 4.7 9.5 14.2
3 Wien 20.0 50.0 70.0 20.0
Austr. total 31.2 18.5 28.3 80.0 6.8
1 Lausanne 68.8 9.3 12.5 90.6 6.3 3.1 9.4

2 Bern 32.1 21.4 32.2 85.7 7.1 7.1 14.2
Swiss total 50.5 15.4 22.5 88.2 3.6 6.7 1.6 11.8
The approximate locations of the Central

European populations involved in this study.
Frequencies of Haplogroups R1b and R1a in Central Europe.
Frequency of Haplogroup I in Central Europe.

52
Frequencies of Haplogroups E3b and J2 in Central Europe.
shows the most frequent haplogroup at various

places in Central Europe. (1) The Iberian Haplogroup R1b forms a west-east
gradient in Central Europe: the percentages are high
Several conclusions can be drawn from the data on the (roughly 50%) in westernmost Germany and low (below
distribution of haplogroup frequencies over the geogra- 20%) in the east (eastern Poland and Hungary).
phy of Central Europe:
(2) The percentages of the Ukrainian Haplogroup R1a
show an opposite tendency: They are low (less than
10%) in westernmost Central Europe and high (more
than 50%) in Poland; they are slightly lower (about
40% or less) in the Czech Republic and Slovakia and
still lower (less than 30%) in Hungary. In the R1b and
R1a maps, the sum of the frequencies of the two haplo-
groups (R1b + R1a) is more or less constant; it is about
70% in Poland, about 60% in the Czech Republic and
Slovakia, and about 40-50% in Austria and Hungary.
(3) The frequencies of the Balkan Haplogroup I in

Central Europe are reflections of two I-centres, the
North-German one (which represents subhaplogroup
I1b2-M223, formerly I1c, and to some extent also the
Scandinavian subhaplogroup I1aM-253) and the West-
Balkan one (which represents subhaplogroup I1b1-P37).
The former is situated within Central Europe and has
values over 30% in northern Germany, while the reflec-
tions of the latter are seen in Central Europe as the
relatively high I frequencies (over 20 %) in Hungary.
The Swiss and Austrian areas with I frequencies less than
10% result from their geographic locations relatively far
Most frequent haplogroups in various
from both I subgroup peak areas in question.
parts of Central Europe. The colours of the four
haplogroups are shown at the lower left corner of (4) The Siberian Haplogroup N3 is rare in Central
the map. The circles and country names with two Europe. Its average frequency is 3.7% in Poland, 3.0%
colours represent cases with equal or almost equal Slovakia, 1.6% in Germany, Austria, and the Czech
frequencies of two haplogroups. Republic, and 0-0.5% in Hungary.
(5) Central Europe has two separate centres for the Early Hungarians of the Great Migration in 500-895 AD.
Farmers’ Haplogroups E+J+G (more precisely those of According to this interpretation, the genome of the
E3b, J2, and G2): The Hungarian centre with frequen- modern Hungarian men is typically Central European
cies of about 20% is a reflection of the E+J+G centre in but their Finno-Ugric language is from the east.
Greece where early farming first arrived from Anatolia
and the Middle East. The Hungarian centre and its
neighbouring areas in Slovakia and the Czech Republic
represent the farmers of the Körös (6000-5500 BCE) Though the Czechs are included in the treatment above,
and Linearbandkeramik (LBK) (4500-3900 BCE) cul- a more detailed analysis is in order. The haplogroups of
tures. The other Central European centre is in Holland. the men of the Czech Republic are analysed thoroughly
This area represents the other main branch of Early in Luca et al. (2005), and the frequencies are summa-
Farmers who expanded from Greece along the Mediter- rized in .
ranean coast to the west and came to Central Europe
along the Atlantic coast through France. The sum total of the frequencies of the most common
three haplogroups cover 74-84% of all haplogroups in
Central European men represent three main types: the Czech Republic. The most common haplogroup is
R1a (about 35%), second is R1b (about 28%), and third
(a) The ”German type” came originally from two main is I (about 18%). The Czech area is homogeneous; only
sources, the Iberian refuge and the Balkan refuge; the two clinal gradients seem to exist: (a) In the southeastern
number of those coming from the Iberian refuge is corner of the country (Brno and Trebic), R1a seems to
slightly higher (about 45%) than those coming from the be somewhat more frequent than elsewhere. This is
Balkan refuge (about 40%); see expected on the basis of the Central European centre of
concerning Haplogroups R1b, I1a and I1b2- R1a in Poland and Slovakia. (b) The frequencies of I are
M223 (labeled as I1c on ). The number of those higher in the west and lower in the east.
coming from the Ukrainian refuge (R1a) is lower (about
5%). The northeastern (Siberian and linguistically Finno-Ug-
ric) Haplogroup N3 has frequencies of about 2-3% in
(b) The “West-Slavic type” came mostly (about 30- the Central areas of the Czech Republic; the average
50%) from the Ukrainian refuge; a smaller portion of total for the entire country is below 2%.
them (about 25%) came from the Balkan refuge and a
still smaller portion (about 15%) from the Iberian ref-
uge; see the R1a, I1a, and I1b1-P37 maps above.
By “North Europe” I mean here the area consisting of
(c) The “Hungarian type” is characterized by the fact the Scandinavian peninsula (Norway and Sweden), Den-
that about equal numbers (about 25-30%) of these men
arrived from the three refuges, the Iberian, Balkan, and
Ukrainian refuges; see the maps concerning R1b, R1a,
I1a, and I1b.
In addition, all three main types of the male populations

in Central Europe received about 4-21% of their men
from the three groups (E, J and G) representing the Early
Farmers from the Middle East. The exact percentage of
the farmers is dependent on the geographic location of
the population in relation to the Middle East: The fre-
quency is highest (21%) in the Hungarians.
As seen in the maps, the three genetic types represent

also three linguistic groups. In this paper, I do not,
however, deal in detail with the possible explanations
for the emergence of the linguistic difference between the
Germanic and Slavic languages. One possibility is the
difference arose from the original linguistic differences
between the Iberian and Balkan refuges, but there are Frequencies of three haplogoups, R1b,
other possible explanations (see, for example, Wiik R1a, and I, in five Czech localities: Klatovy (K),
2002). The Hungarian language (unlike the majority of Pisek (P), J.Hradec (H), Trebic (T), and Brno (B).
the genetically defined modern Hungarian people) ar- The relative positions of the figures for the three
rived from the southern Ural area with the ancient haplogroups are shown in the lower left corner.
54
mark, Iceland, Finland, and Karelia. The continental sense that about 96-98% of its Y-chromosomes are
part of this area is often called Fenno-Scandia. members of just four haplogroups: R1b, R1a, I, and N3.
Each of these is from a different Ice-Age refuge, which
Linguistically, the people of this area represent two means that the male populations of North Europe came
language phyla, Indo-European and Finno-Ugric. The originally from the Iberian, Balkan, Ukrainian, and
IE languages are represented by the Germanic, more Siberian refuges. Therefore, they represent the Old
precisely the North Germanic or Scandinavian langua- Europeans (not the Early Farmers). The percentages of
ges, and the FU languages by two main branches of the the individual haplogroups are, however, quite different
“Early Proto-Finnic” languages, more precisely two in various parts of North Europe, which makes it possi-
Finnic languages (Finnish and Karelian) and Saami. The ble to draw conclusions about where the North Europe-
four Scandinavian languages, Icelandic, Danish, Norwe- ans originally came from. The overall frequencies of the
gian, and Swedish are spoken in their respective coun- haplogroups in the seven North-European male popula-
tries. Swedish is spoken, in addition, on the Borthian tions are seen . Next, the main four haplogroups
and Newland (Uusimaa) coasts of Finland as well as in of Northern Europe will be considered separately.
the archipelago between Finland and Sweden. Finnish is
spoken outside Finland in northern Sweden and Nor-
way. The Karelian language is spoken only by a minor-
ity in the Russian Republic of Karelia. The frequencies of the “Iberian” Haplogroup R1b
(generally thought to be the oldest in Europe) are seen in
Genetically the North European male population (like . The group is typically West European; its
many other European populations) is concise in the highest frequencies (about 80-90%) are found in Ireland
1 43.7 22.4 33.1 0.3 99.5 0 0 0 0 0 0

2 43.2 13.7 34.2 0.5 91.6 4.3 3.2 0.2 7.7 0.4 0
3 27.5 24.6 41.4 3.9 97.4 2.2 1.3 0 3.5 0 0
4 22.4 17.0 47.9 8.2 95.5 1.6 1.4 0.4 3.4 0.9 0.6
34.2 19.4 39.2 3.2 96.0 2.0 1.5 0.2 3.7 0.3 0.2
5 6.8 14.2 33.8 42.6 97.4 0 0 0 0 0 0

6 1.4 8.9 25.7 61.2 97.2 0.5 0 0 0.5 1.3 1.0
7 1.3 40.7 20.5 38.0 100 0 0 0 0 0 0
3.2 21.3 26.7 47.3 98.4 0.2 0 0 0.2 1.4 0.3
* NG = North German Language Group; FU = Finno-Ugric Language Group.
17.5 12.5 45.0 5.0 10.0 90.0 1.8 0 0 0 0 5.0 5.0 40

25.4 12.7 38.2 0 14.6 90.9 0 5.5 0 5.5 0 1.8 1.8 55
31.7 14.6 29.3 2.4 7.3 85.4 2.4 9.8 0 12.2 0 2.4 0 41
21.9 17.1 41.5 4.9 2.4 87.8 0 0.0 4.9 4.9 2.4 2.4 2.4 41
40.0 8.9 35.6 2.2 2.2 88.9 2.2 4.4 0 4.4 0 6.7 0 45
16.7 14.3 45.2 4.8 9.5 90.5 0 4.8 2.4 7.1 0 2.4 0 42
17.1 9.8 24.4 17.1 19.5 87.9 2.4 0 4.9 4.9 2.4 2.4 2.4 41
24.6 11.8 37.0 4.9 9.5 88.5 1.2 3.6 1.6 6.4 0.7 3.3 1.6 305
F = F*(xG,I,J,K), K = K*(xN3,P), P = P*(xR1a,R1b3).
26.8 27.1 34.7 10.6 0.8

27.1 31.5 39.7 ~ 1-2
43.2 24.3 30.2 ~ 1-2
26.4 26.8 41.4 3.2 2.2
43.2 24.3 30.2 2.3
35.5 28.0 33.3 ~2 ~1
35.8 19.3 40.4 ~4
44.7 13.2 42.1
35.3 24.3 36.5 2.9 1.0
0 8.3 16.7 2.0 70.8 0 2.1 48

4.5 4.5 18.2 0 68.2 0 4.5 22
0.9 4.7 14.9 2.0 78.5 0 0 107
4.6 6.2 23.3 0 65.1 0 0.8 129
2.6 5.9 18.9 0.6 70.9 0 1.9 306
3.4 19.0 46.6 0 25.9 3.4 1.7 58

8.0 12.0 36.0 0 40.0 0 4.0 25
6.2 8.3 52.1 2.0 27.1 0 4.2 48

6.1 2.0 34.7 0 55.1 0 2.0 49
4.0 2.0 28.0 2.0 60.0 0 4.0 50
5.2 8.7 40.0 0.9 41.3 0.9 3.2 230
3.7 7.1 28.0 0.7 58.2 0.4 2.5 536
and on the whole in the western parts of the British Isles. of the Recolonization of Northern Europe that started
In Germany, its percentage is slightly less than 50%, and after the Late Glacial Maximum and has continued in
about the same percentages (43-45%) are also found in many phases after that.
the westernmost areas of North Europe, Denmark,
western Norway and Iceland. The other extreme in
North Europe is represented by the Finns and Saami
whose R1b percentages are only 0-8% (average about The “Ukrainian” Haplogroup R1a shows, somewhat
4%). Between the two extremes, there are three inter- surprisingly, a west-east gradient in North Europe: The
mediate zones in the map: (a) In southwestern Scandina- frequency is (a) highest (32%) in west-central Norway,
via, the percentages are 32-38%, (b) in northern (b) slightly lower (about 24-28%) in many other parts
Norway and southeastern Sweden 22-27%, and (c) in of Norway and in Iceland, (c) about 10-19% in southern
northern Sweden and Gotland 15-17%. Thus, the R1b and northern Norway, Denmark, Sweden, and Finnish
percentages form a west-east gradient according to Bothnia, and (d) only about 2-8% in the other parts of
which the percentage descends from about 45% to zero Finland. This frequency distribution may seem surpris-
from Denmark and Southern Norway to Eastern Finland. ing because the European peak area (frequency about
55%) is in Poland. The frequency distribution makes
The gradient is a reflection of the migrations from West one believe that there has been a movement of R1a men
Europe (the Atlantic Coast) and ultimately from the from Central Europe to the Central-Norwegian coast.
Iberian refuge to Scandinavia. The migrations were part This expansion represents the Ahrensburgian culture
56
Frequencies of Haplogroups R1b and R1a in North Europe and its vicinity. The North
European parts of the maps are based on Tables 4, 5, and 6.
Frequencies of Haplogroups I and N3 in North Europe and its vicinity. The North
European parts of the maps are based on .
(about 8500 BC), perhaps also the Hamburgian culture in Belarus. In Russia, there is a south-north gradient (cf.
(about 15-13.7 kya) of the northern parts of Central ).
Europe (cf. Saukkonen 2006, p. 72). The route of this
expansion was western (through Denmark and southern Linguistically, the N3 men are generally thought to
Scandinavia) rather than eastern (through Balticum and represent the speakers of Finno-Ugric languages (in
Finland): this is shown by the fact that R1a-frequencies western Siberia; they represent the speakers of
are low in Finland. As a matter of fact, Finland is very Altaic/Turkic languages).
much like a vacuum in this respect in North Europe: In
Karelia and the Baltic countries (Estonia, Latvia, and The Haplogroups E3b, J2 and G of the Early Farmers,
Lithuania), R1a-frequencies are of the order of 35-42 occur in Scandinavia, while among the Finns, Karelians,
(i.e. even higher than on the central Norwegian coast). and Saami, these haplogroups are practically non-exis-
tent. The sum total of the frequencies of these Haplo-
groups (E3b+J2+G) is highest (4.3+3.2+0.2 = 7.7%)
among the Danes and lower among the Norwegians
Haplogroup I is generally thought to have spread to its (2.2+1.3+0= 3.5%) and Swedes (1.6+1.4+0.4 = 3.4%).
modern areas from the Balkan refuge. The haplogroup The Icelanders do not have these haplogroups.
has many subgroups. The most frequent of these in
North Europe is I1a; Haplogroup I1b2-M223 (formerly
I1c) is common in North Germany and Haplogroup
I1b1-P37 (formerly I1b) in the western Balkans. As seen By “East Europe” is meant here the geographic area
in , Haplogroup I (most of which consists of covered by most of the European parts of Russia, the
Haplogroup I1a-M253 in North Europe) is common Ukraine, Belarus, Romania, Moldovia, Lithuania, Lat-
over almost all of North Europe: frequencies of 30-50% via, and Estonia. The area is linguistically heteroge-
are found almost everywhere in North Europe; the only neous in that it includes languages of four language
exception is eastern Finland (and evidently also Karelia) phyla: (1) Indo-European, (2) Finno-Ugric (Uralic), (3)
where the frequencies are below 20%; there is a rather Altaic, and (4) North Caucasian. The Indo-European
steep west-east gradient in Finland, the frequencies being languages belong to East Slavic, Romance and Iranian
about 50-40% in the west and below 20% in the east. groups. The individual East Slavic languages are Rus-
As seen from the frequencies of Germany (25%) and sian, Belarussian, and Ukrainian. The Finno-Ugric
Poland (17%), as well as those in East Europe (7-19%), (Uralic) language groups are Finnic (e.g. Estonian), Vol-
North Europe forms an independent island of Haplo- gaic (Mordvian and Mari), Permic (Udmurtian and
group I. Komi), Ugric (Hungarian) and Samoyed (Nenets). The
Altaic languages belong to the Turkic group (Turkish,
In Finland, Satakunta is exceptional in having a frequen- Tatar, Chuvash, and Bashkirian). The North Caucasian
cy as high as 52% for Haplogroup I. In Sweden also, languages are represented, for example, by Chechenian.
there is one rather exceptional area: The “German”
Haplogroup I1b2-M223 is as high as about 14% in
Västerbotten. The average total of this haplogroup is
below 5% in Sweden as a whole. The frequency of Haplogroup R1b is very low (below
10%) in Russia, but it rises to about 20-40% in some
parts of the Caucasus (cf. the Wikipedia and Balanovsky
maps and ). In East Europe is seen to
As seen in , Haplogroup N3 is typically eastern. consist of three separate areas (cf. the three shades of
Its total area extends as far east as the Pacific Ocean, and gray):
it has very high frequencies (85%) in the Yakuts in
northeastern Siberia. In North Europe, Haplogroup N3 (a) The southeastern corner (Baskirs and Ossetians) has
is commonest in Eastern Finland (71-78%). The per- R1b values (43-47%) of the “West European” type.
centage diminishes with geographic distance outside The explanation is that a considerable portion of the
Eastern Finland and the percentages are lower (53-68%) R1b or R1 men first arriving in eastern Europe from
elsewhere in Finland. The percentage is slightly lower in Asia about 40 kya stayed in the steppe and mountainous
the East-Karelians and Vepsians (38%), as well as in the areas around the Caspian Sea and the Caucasus.
Estonians, Latvians, and Lithuanians (34-42%). In
Scandinavia, there is a northeast-southwest gradient, the (b) The intermediate zone with R1b frequencies of 10-
N3-percentages being about 10-15% in the northeastern 19% is situated in the southern and eastern parts of East
and western Scandinavia and very low in the south and Europe and consists of the following populations:
west (in Southern Norway it reaches zero). To the south Nenets (19%), Komi (16%), Chuvash (12%), Mordvi-
of North Europe the N3-percentage is very low: accord- ans (13%), Ukrainians (11%), Belarussians (10%), Lat-
ing to the map, 2% in Germany, 3% in Poland, and 4%
58
7.9 31.9 20.1 33.9 0 93.8 1.8 0.5 0.7 3.0 3.5
11.8 40.5 8.4 37.8 98.5 0.2 0 0.2 0.2?
4.5 38.3 13.3 42.2 98.3 1.2 0 0 1.2 0.2
9.9 40.0 29.7 2.8 82.4 6.7 3.3 1.3 11.3 1.5
10.7 45.4 16.0 7.6 79.7 3.1 6.3 9.4 9.8?
13.0 20.4 46.3 0 0 79.7 7.4 5.7 5.6 18.7

16.7 27.6 30.9 1.9 0.7 77.8 12.8 7.7 1.0 21.5 1.0 1.0
12.5 19.7 27.9 2.1 0 62.2 13.3 5.9 13.8 33.0 3.2
4.0 32.0 36.0 29.0 21.0 50.0 7.0 8.0

13.0 13.0 3.0 74.0 77.0 10.0
5.4 34.2 13.1 35.5 7.5 95.7 0.2 1.8 1.2 3.2 0 1.5
7.5 46.5 15.3 16.3 0.5 86.1 5.0 3.4 0 8.4 1.7 1.7
4.8 55.4 21.0 9.5 0.5 91.2 1.8 3.5 1.0 6.3 1.4 1.1
13.3 26.5 19.2 16.9 2.4 78.3 ? ? ? ? ? ? ?

5.0 21.0 4.2 51.3 0 81.5 6.3 6.3 11.8
9.2 30.5 3.5 36.6 14.4 94.2 2.4 2.4 4.8 2.3
16.0 23.7 4.5 36.2 24.3 104? 0 0 0 0
19.1 4.2 38.2 38.2 99.7 0
0 0 0 51.7 44.9 96.6 0 0 0 0 0 0 3.4
0 0 0 23.7 74.6 98.3 0 0 0 0 0 0 1.7
6 29 16 25 0 76.0 2 8 10.0 7.0 6.0

12 18 24 18 0 72.0 6 6 12.0 18.0
47 26 ? 17 0 90 ? ? ? ? ? 10?
vians (12%), Moldovians (17%), Gagauzes (13%), and mixed populations in the sense that about half of their
Romanians (13%). men represent the ancient mammoth hunters from the
Siberian refuge (representing Haplogroup N3) and an-
(c) The R1b values are very low (1-9%) in the zone to other half from the Ukrainian refuge (representing Hap-
the northwest of the intermediate zone. This zone con- logroup R1a).
sists of the following populations: Northern, Central
and Southern Russians (5%, 8% and 5% respectively), According to the Wikipedia and Balanovsky R1a maps,
Udmurts (9%), Mari (5%), Lithuanians (3%), and Esto- the European maximum area (with frequencies over
nians (8%), as well as the North European populations 50%) of Haplogroup R1a is in Poland, and the area of
Finns (1%) and Saami (7%)). almost equally high frequencies (over 40%) extends to
Belarus. In East Europe, there is a west-east gradient
and the frequencies descend to about 20-30% in north-
ern Russia and close to zero in the southeastern corner
The frequencies of Haplogroup R1a are high (about (the Caucasus) of Europe.
30-50 %) in the following East-European populations:
Estonians, Latvians, Lithuanians, Belarussians, Ukraini- According to , the “Polish” maximum area
ans, Russians, and Tatars. These are evidently the actually extends to southern Russia (55%) and there is
populations that (at least partly) used the Ukrainian a zone with frequencies over 40% in practicality all the
refuge during the Ice Age. This means that the Esto- eastern parts of East Europe. This zone contains in
nians, Latvians, Lithuanians, and North Russians are addition to the North European Karelians (41%), the
Frequencies of Haplogroups R1b and R1a in East Europe. The maps are based on .
Frequencies of Haplogroups I and N3 in East Europe. The maps are based on .

60
following East European populations: Latvians (41%), rare in the easternmost populations: the frequency is 5%
Lithuanians (42%), Belarussians (40%), Central Rus- in the Komi and 4% in the Udmurts.
sians (47%), and Ukrainians (45%). The next zone,
with frequencies of 30-39% consists of the Estonians Regarding the I frequencies of the East Europeans, one
(32%), Northern Russians (34%), and Udmurtians should keep in mind that these populations have not
(31%). Still further from the peak area, with frequencies received their Haplogroup I men only from the I1b1-P37
of 20-29%, are the populations of the Komi (24%), maxima in the Balkans and Romania. Particularly in the
Bashkirs (26%), Tatars (29%), Mari (21%), Mordvians northwestern parts (e.g. in Estonia and Latvia), a major-
(27%), Moldovians (28%), Gagauzes (20%), and Ro- ity of the representatives of Haplogroup I have come
manians (20%). The R1a frequency is somewhat lower from the peak areas of Haplogroups I1a and I1b2 in
(10-19 %) in the Chuvash population, and still lower southern Scandinavia and Germany. So, for example,
(4%) in the Nenets in the northeastern corner of Europe. about 82% of the Estonian men of Haplogroup I belong
Also the Finns (considered here to represent North, to the “Scandinavian” subgroup I1a and only about 2%
rather than East Europeans) have a very low R1a fre- of them belong to the “German” Haplogroup I1b2-
quency (9%). The low value of the Finns tends to show M223; about 16% of the Estonian men belong to the
that the route of the R1a men of Norway (25% of “Balkan” Haplogroup I1b1-P37; see . These
Norwegians in the map) followed the western route figures tend to show that about 82+2 = 84% of the
from Poland to Denmark and finally to Norway, and Estonian men of this haplogroup came from the west
not to the same extent the eastern route from East and only about 16% from the south. The equivalent
Karelia through Lapland to Norway. Supporting this figures for the Latvians are almost identical: 67+17 =
hypothesis is the frequency for the Saami, which is 84% from the west and 16% from the south. Quite
relatively low at 7%. different ratios are shown by the Romanians: about 8%
of the Romanian men of Haplogroup I represent the
All of the maps for Haplogroup I and its subgroups

show that the most common subgroup of Haplogroup I
in the Balkans is Haplogroup I1b1-P37, which has a
maximum frequency of about 40% and even more in
some parts of the Western Balkans. This subhaplogroup
has a secondary maximum in Romania (maximum fre-
quency about 25-30%). It is particularly the Romanian
maximum that (according to the maps used) has influ-
enced the East European populations. There is in East
Europe a south-north gradient (from about 24-30% to
4%) extending from Romania to the Nenets territory in
the northeastern corner of Europe.
14.8 2.9 0.5 18.2
The other two subhaplogroups of I are less frequent in 4.7 1.1 1.2 7.0
East Europe. The maximum area of I1a-M253 is in
southern Scandinavia and that of I1b2-M223 is in
Germany. Both have, however, a secondary maximum 55.7 0 3.6 59.3
in the eastern parts of East Europe. I1a reaches as high
values as 15% in the Volga area, and I1b2 reaches the 26.8 0 0 26.3
frequency of 6% in the Ural area; see the Wikipedia and 38.9 0 1.4 40.3
Balanovsky I1a maps.
40.0 0.4 0.9 41.3
In all three subhaplogroups of I are put togeth- 19.0 0 0.7 19.6
er. It shows the Romanian maximum area (46%) and
outside it the zone of about 30% in Moldovia (31%) 28.6 0 0 28.6
and Belarus (30%). The next zone with frequencies of 25.0 0 12.5 37.5
20-29% is further away from Romania: Moldovians
(28%), southern Russians (21%), Chuvash (24%), and 16.7 0 10.0 26.7
Estonians (20%); also the Karelians (21%) and Finns
(26%) in North Europe belong to this zone. The next
1.7 17.7 1.9 21.3
zone has frequencies 10-19%: Ukrainians (16%), Osse-
tians (11%), Mordvians (19%), Central Russians 3.3 21.7 3.3 28.3
(15%), and Northern Russians (13%). Haplogroup I is
2.5 24.1 5.1 31.7
gives the frequencies of the commonest three original FU language to an IE (Baltic and Slavic) one.
subhaplogroups of I in eleven European areas (Rootsi et The language shift took place particularly at the time of
al. 2004a). The figures show, for example, that a the arrival of agriculture and the “Slavic Expansion”
majority of the Estonian and Latvian men of Haplo- more than a thousand years ago.
group I represent the “Scandinavian” Haplogroup I1a
and only a small minority the “Balkan-Romanian” Hap- The peak area of the frequencies of the “Siberian”
logroup I1b1-P37. Haplogroup N3 in Europe is in Eastern Finland (70% in
the N3 Maps above). In , there is a secondary
The men of Haplogroup I have clearly arrived in the maximum (51%) in the Mari area. Many of the north-
Baltic area from two directions: from the west and from ern populations of East Europe have frequencies of
the south. The former men represent “Germanic” (i.e. 34-39%: Lithuanians 39%, Latvians 38%, Estonians
“German” and “Scandinavian”) subhaplogroups I1a- 34%, Northern Russians 36%, Udmurts 37%, Komi
M253 and I1b2-M223 and the latter those of East-Euro- 36%, and Nenets 38%. In the next zone to the south
pean subhaplogroup I1b1-P37. The two directions of and east of the Volga area, the percentages diminish
steeply: Tatars 25%, Central Russians 16%, Mordvians
17%, Chuvash 18%, Bashkirs 17%, and Southern Rus-
sians 10%. In the next, more southern zone, the fre-
quencies are even lower: Belarussians 3%, Ukrainians
8%, Moldovians 2%, and Gagauzes 2%. As seen from
the map, the frequencies of Haplogroup N3 in East
Europe form a regular north-south gradient.
(5) Haplogroups E3b, J2, and G belonged to the first

farmers to arrive in Europe. E3b is high in the Moldo-
vians, Turks and Gagauzes; J2 is high in some Caucasian
populations like Ossetians as well as Turks; and G is
high (in addition to some Caucasus populations like
Ossetians) also in the Romanians and Gagauzes. Two
haplogroups of the ancient farmers, E3b and J2, show
clear south-north gradient: the frequencies of E3b and
J2 are about 15 and 10% in Romania and they descend
gradually through the Ukraine and Russia and approach
zero in the northeastern parts of Europe.
Frequencies of Haplogroup I in some East

European populations. Lithuania is considered as six
separate subareas in the map. Source: Kasperavièiûtë et Next three East-European populations are treated sepa-
al. (2004). rately and in more detail by three language groups: (a)
Russians, (b) Turkic-speakers, and (c) Balts and Finno-
Ugrians.
arrival are seen also from the Lithuanian I-frequencies
shown in , in which the highest frequencies
(16.9% and 17.5%) are in the westernmost and eastern-
most areas of Lithuania, while in the intermediate areas
the frequencies are only 2.9-9.4%. A similar two-peak
frequency distribution (high in the west and east and low
in the middle) is expected in Latvia.
(4) The following eight populations form a homogenous

group on the basis of high (30-50%) N3 values: Estoni-
ans, Latvians, Lithuanians, North Russians, Mari, Ud-
murts, Komi, and Hanti. The ancestors of these
populations can be called the ancient “mammoth hunt-
ers” of Northeastern Europe. The populations were a
genetically and probably also linguistically homogene-
ous population. The language was, supposedly, Finno- The three main dialects of European
Ugric (Uralic). This, however, implies language shift: Russian: North Russian, Central Russian, and
the Baltic and north Russian populations shifted their South Russian.
62
A. Russians
The Russians are treated above as three separate popu-

lations: North Russians, Central Russians, and South
Russians. This division reflects the three main dialects
of Russian, as shown in .
Geographically more detailed data on the frequencies of

the Y-chromosome haplogroups in traditional Russia
are given in . The approximate locations of the
fourteen localities of the table are shown on .
The table is a simplification of the corresponding table
in Balanovsky et al. (2008).
The frequency zones of the eight haplogroups of

are seen in . The maps allow the following
generalizations about the Russians:
(1) The Russians consist mainly of two populations: (a)

almost half (about 45%) of them represent the
“Ukrainian” Haplogroup R1a, and (b) about one quar-
ter of them represent the “Siberian” Haplogroup N
(more precisely N3 + N2). The former are more fre-
The locations (shown as dots) of the quent in Southern Russia and the latter in Northern
fourteen russian populations analysed in Russia. Haplogroup R1a forms a south-north gradient,
Balanovsky et al. (2008). and, conversely, Haplogroup N3 forms a north-south
0 44.4 0 0 46.3 7.4 98.1 0 0 0 1.9 54

14.0 39.5 0.9 4.4 23.7 15.8 98.3 0 0 0 1.7 114
6.6 19.8 12.1 9.9 36.3 3.3 88.0 0 5.5 5.5 6.5 91
0.8 33.1 11.6 8.3 35.5 3.3 92.6 0.8 0.8 1.6 5.8 121
5.4 34.2 6.2 5.7 35.5 7.5 94.3 0.2 1.6 1.8 4.7 380
11.5 32.7 11.5 11.5 13.5 0 80.7 5.8 3.8 9.6 9.7 52
6.8 56.2 2.7 8.2 11.0 0 84.9 4.1 4.1 8.2 6.9 73
5.3 52.6 3.5 10.5 15.8 1.8 89.5 3.6 0 3.6 6.9 57
2.7 45.3 6.7 9.3 28.0 0 92.0 4.0 1.3 5.3 2.7 75
11.2 45.8 1.9 10.3 13.1 0.9 83.2 7.5 2.8 10.3 6.5 107
7.5 46.5 5.3 10.0 16.3 0.5 86.1 5.0 2.4 7.4 6.5 364
3.6 62.7 8.2 13.6 4.5 0.9 93.5 0.9 0.9 1.8 4.7 110
2.2 55.6 4.4 17.8 13.3 0 93.3 2.2 2.2 4.4 2.3 45
5.2 59.4 3.1 16.7 6.3 0 90.7 1.0 1.0 2.0 7.3 96
2.8 59.4 3.5 12.6 11.9 0.7 90.9 0.7 4.2 4.9 4.2 143
8.8 47.3 4.4 16.5 6.6 1.1 84.7 3.3 4.4 7.7 7.6 90
4.8 55.4 3.9 15.9 9.5 0.5 90.6 1.8 3.0 4.2 5.2 484
Frequencies of Haplogroup R1a in Frequencies of Haplogroup R1b in

historical Russia. historical Russia.
Frequencies of Haplogroup N3 in Frequencies of Haplogroup N2 in

64
Frequencies of Haplogroup I1a- Frequencies of Haplogroup I1b1-

M253 in historical Russia. P37 in historical Russia.
Frequencies of Haplogroup J2 in Frequencies of Haplogroup E3b in

gradient. The language of the southern areas has been sia, these haplogroups do not exist (the explanation
typically Indo-European, while that of the northern being, of course, that these areas were not suitable for
areas has been Finno-Ugric. The boundary between the early farming because of their cold climate and the acid
two has been moving from the south to the north, soil of the conifer forest).
initially as a result of the arrival of agriculture, and later
as a result of the spread of the Orthodox Church and B. Turkic-Speaking Populations
southern trade. Since the beginning of the Soviet period,
the Russian language has expanded at the expense of the The main Turkic-speaking populations in East Europe
Finno-Ugric languages as a result of systematic political are the Tatars, Chuvash and Bashkirs. shows
policy. Consequently, the area of the Finno-Ugric- the frequencies of these and, in addition, that of the
speaking population (typically hunters rather than farm- Turks (belonging in this study to “The Balkans”).
ers) has diminished and that of the Indo-European (more
precisely East-Slavic) language has expanded. The line shows that the European Turkic-speaking
of language shift from Finno-Ugric to East-Slavic, with populations is not homogeneous. Each of the five
its bilingual intermediate zone, has gradually moved groups (R1b, R1a, I, N3, and J+E3b+G) are represented
towards the north. This process continues even today. differently in the four populations, and each of the four
populations has its own peculiarities:
(2) There are four frequency zones of R1b in Russia.
The frequencies in the zones from the west to the east are (1) The traditional “Iberian” Haplogroup R1b is partic-
as follows: 0-5%, 11-14%, 1-5%, and 7-9%. The ularly high (almost 50%) in the Bashkirs, and much
complicated variation can be explained by the geogra- lower (about 6-16%) in the other Turkic-speaking pop-
phic location of the zone from the West-European and ulations. In this respect, the Bashkirs come close to
East-European centres of R1b: (a) The frequencies are some Caucasian populations, whose R1b-percentage is
highest (11-14%) in the middle zone that has received its almost equally high (about 43%). The high R1b-percent
R1b-men from both centres. (b) The lowest frequencies in the two populations in question is interpreted here
(0-5% and 1-5%) are in areas that are far from both rather as an “Asian” or “Caucasian” (than “Iberian” or
centres. (c) The frequencies are second highest (7-9%) “West-European”) feature.
in the zone that is relatively close to the East-European
centre of R1b. (2) The “Ukrainian” Haplogroup R1a is relatively high
(18-29%) in the Turkic-speaking populations (Tatars,
(3) In the northeastern corner of historical Russia, the Chuvash, and Bashkirs) of the Volga and Ural areas, but
two subhaplogroups N3 and N2 are, to some extent, considerably lower (about 6%) in the Turks of the
complementary: in the area where the frequencies of N2 Balkans and Anatolia.
are relative high, those of N3 are relatively low and vice
versa. (3) The “Balkan” Haplogroup I represents the opposite
of Haplogroup R1b: its frequency is low (less than 5%)
(4) I1b1-P37 is common (13-18%) in Southern Russia, in the Bashkirs but higher (14-24%) in the other Turkic-
but is almost non-existent (0-4%) in northernmost Rus- speakers.
sia.
(4) The “Siberian” Haplogroup N3 varies from about
(5) The “Scandinavian” Haplogroup I1a is common 25% in the Tatars through about 17-18% in the Chuvas
(about 8-12%) in Central Russia. One natural explana- and Baskirs to only a few percent in the Turks.
tion is the Vikings.
(5) Haplogroups E3b, J2 and G2 of the Early Farmers
(6) The Early Farmers’ Haplogroups J2 and E3b are are high (about 38+12+3 = 53%) in the Turks, but much
most common (3-6%) in Central Russia and slightly less lower (close to 10%) in the other three populations.
common in (1-4%) Southern Russia; in Northern Rus-
6 29 16 25 76 2 8 0 10 4 7 2
12 18 24 18 72 6 6 0 12 18
47 26 <5? 17 ~92 <5? <5? <5? ? ? ? ? ?
13.3 5.8 14.2 2.2 35.5 11.7 38.2 3.3 53.2 1.7 9.9
19.6 19.7 ~14 15.6 ~69
66
C. The Balts and Finno-Ugrians Tundra Nenets, (b) about equally high (about 38%) in
the Hanti of Northwestern Siberia, (c) lower (about
The last language group of the East-European popula- 14-24%) in the Komi and Udmurts, and (d) non-existent
tions to be dealt with consists of two linguistic sub- or almost non-existent in the other FU-speaking popula-
groups. One subgroup speaks Baltic languages and the tions. (In this respect the Vepsians who are close rela-
other, Finno-Ugric (Uralic) languages. The treatment of tives of the Karelians are an exception: the N2-frequency
the two language groups as one is based on the fact that of the Vepsians is as high as about 17% (Rootsi et al.
the Balts and Estonians are genetically close to each 2006).
other.
(5) The frequencies of the three haplogroups of the Early
summarizes the haplogroup frequencies for Farmers (J+E3b+G) are low (about 2-6%) in all the
these eight populations, and allows the following gener- FU-speaking populations. This results, of course, from
alizations to be made: the northern habitats of these populations.
(1) The frequencies of the “Iberian” and

“Asian/Caucasian” Haplogroup R1b are 5-16%. Part
of the men of this haplogroup are from the west and part The region defined here as the home of the “Balkan
from the east. Europeans” is shown on . The Balkans are often
(2) The frequencies of the “Ukrainian” Haplogroup R1a

are about 20-40% in almost all the European FU-speak-
ing populations. The only exception are the Tundra
Nenets who do not have this haplogroup. (R1a is also
very low or non-existent in the FU-speaking populations
(e.g. Hanti) of Siberia.)
(3) The frequencies of Haplogroup I form two catego-

ries: Estonians and Mordvians belong to one category
with frequencies of about 20% and the Komi, Udmurts
and Mari to the other with frequencies of about 4%.
(4) With respect to the frequency of the “Siberian”

Haplogroup N3, the Finno-Ugric speaking populations
form two categories: this haplogroup is relatively low
(about 17%) in the Mordvians, but high (34-51%) in
the other populations. The feature makes the Mordvi-
ans differ from their linguistic relatives in the direction
of more southern populations.
On the basis of the “Samoyedic” Haplogroup N2, the

FU (or Uralic) speakers form four categories: N2-fre- The Balkan region (south of the dark line).
quency is (a) highest (about 45%) in the (Samoyedic)
R1b R1a I N3 N2 OE E3b DE J G EF K other

11.8 40.5 8.4 37.8 98.5 0.2 0 0.2 0.2?
4.5 38.3 13.3 42.2 98.3 1.2 0 0 1.2 0.2
7.9 31.9 20.1 33.9 0 93.8 1.8 0.5 0.7 3.0 3.5
13.3 26.5 19.2 16.9 2.4 78.3 ? ? ? ? ? ?
5.0 21.0 4.2 51.3 0 81.5 6.3 6.3 11.8
9.2 30.5 3.5 36.6 14.4 94.2 2.4 2.4 4.8 2.3
16.0 23.7 4.5 36.2 24.3 >100? 0 0 0
0 0 0 23.7 74.6 98.3 0 0 0 0 0 0 1.7
Eighteen localities in the Balkans: 1 = Slovenes, 2 = Croats, 3 = Bosniacs (Bosnia-Herzegovians),

4 = Serbs, 5 = Bulgarians, 6 = Macedonians, 7 = Albanians, 8-17 = Greeks, and 18 = Turks.
defined geographically by the Danube-Sava-Kupa line The Balkan populations represent four language groups:
(the black line on ), according to which Slovenia, (1) South Slavic, (2) Albanian, (3) Greek, and (4) Turk-
northern Croatia, and northern Serbia do not belong to ish. The first three represent IE languages; Turkish is an
the Balkans, but the eastern coast of Romania does Altaic language. The three IE languages or language
belong to it. However, in this study, Slovenia (but not groups and Romanian form a special group often called
Romania or Moldovia), entire Croatia, and entire Serbia the “ ”. The languages share little
are included in the definition. common vocabulary but they show great similarity in
grammar; so, for example, they have very similar case
systems and they all have become more analytic. These
features can be interpreted as an indication of language
shifts having taken place in the area.
The most frequent haplogroup in the Balkans is Haplo-

group I, or more precisely, its subhaplogroup I1b1-P37.
The fact that R1a and R1b are almost equally frequent
in the Balkans indicates that the Balkans lie at the
west-east dividing line of Europe or slightly on its east-
ern side. The high E3b anf J2 haplogroup values indi-
cate that the Balkans belong to an area of early farming.
The following gradients can be found in the Balkan area:
(1) There is a north-south gradient of R1b in the north

of the Balkans: The frequency of R1b is about 25% in
the Slovenes, but is considerably lower elsewhere and
only about 2% in some parts of Greece. The gradient is
a reflection of the spreading of the West-European or
Iberian men to the Balkans (the other alternative expla-
The frequencies of Y Haplogroup I in nation is that these men have come to the Balkans
eighteen Balkan populations. A large majority of directly from the Caucasian/Asian R1b-areas).
the haplogroup represents subhaplogroup I1b1-
P37. Italian averages (in all the Balkan maps) are (2) Haplogroup R1b exhibits a north-south gradient,
based on the frequencies of four eastern localities with the frequencies of this haplogroup at 10-15% in the
in central Italy: Pescara, Foggia, North Gargano, northern part of the Balkans and only about 3% in the
and Brindisi (Di Giacomo et al. 2003). southern part.
68
25.5 29.5 30.4 0 85.4 12.0 4.5 0.5 17.0 0 0

2.2 12.2 73.3 0 87.7 8.9 1.1 1.1 11.1 1.1 0 90
15.7 34.3 37.0 87.0 5.6 1.9 0.9 8.4 1.9 1.9 1.9 108
16.4 38.4 27.3 82.1 6.8 10.9 17.7 73
6.4 27.7 53.2 87.3 4.3 6.4 10.7 2.1 2.1 47
6.4 8.8 65.9 81.1 4.4 3.3 1.1 8.8 1.1 14.3 1.1 91
1.1 20.5 53.0 74.6 3.8 2.3 10.6 16.7 1.5 6.1 1.5 132
3.5 15.3 48.2 0 67.0 12.9 11.9 3.5 28.3 3.5 1.2 85
6.2 13.6 35.8 0 55.6 22.3 8.7 1.2 32.2 4.9 7.4 81
17.0 12.0 42.0 0 71.0 17.0 12.0 0 29.0 0 0
10.0 35.0 20.0 0 65.0 15.0 20.0 0 35.0 0 0
17.6 9.8 19.6 0 47.0 21.6 27.5 2.0 51.1 0 2.0
13.4 11.0 16.0 0 40.4 22.7 21.3 5.9 49.9 7.8 0.4 366
19.0 4.8 23.8 0 47.6 9.5 28.6 4.8 42.9 9.5 0 21
16.7 8.3 8.3 0 33.3 29.2 20.9 4.2 54.3 12.5 0 24
11.1 5.6 11.1 0 27.8 44.4 16.7 0 61.1 11.1 0 18
8.0 20.0 12.0 0 40.0 28.0 16.0 12.0 56.0 4.0 0 25
12.0 8.0 36.0 0 56.0 24.0 16.0 4.0 44.0 0 0 25
5.0 25.0 20.0 0 50.0 20.0 15.0 5.0 40.0 10.0 0 20
19.0 9.5 14.3 0 42.8 14.3 28.6 4.8 47.7 9.5 0 21
9.8 7.7 14.0 0 31.5 15.4 38.5 7.0 60.9 7.7 0 143
7.4 11.1 18.5 0 37.0 18.5 29.6 7.4 55.5 3.7 3.7 27
26.2 9.5 2.4 0 38.1 23.8 19.1 9.5 52.4 9.5 0 42
13.3 5.8 14.2 2.2 35.5 11.7 38.2 3.3 53.2 1.7 9.9
12.1 16.0 33.3 0.2 57.1 16.0 16.1 1.9 36.8 0.7 0.9 0.7 3.6 0.7
Y* = Y (x A, DE, G2, I, J, P) = {e.g., K2, L, NO, GxG2, H}
(3) The frequency of Haplogroup I1b1-P37 is very high 25.5% for the Slovenes and 20% for the Hungarians; in
(about 40%) in the Western Balkans and it diminishes in the other Balkan populations, the R1b values are consid-
all directions, becoming as low as about 10-5% in the erably lower (11-17.6%). This is an example of geo-
northernmost and southernmost parts of the Balkans; graphic nearness being a more relevant factor than
there is, however another secondary centre with values linguistic relatedness. The Slovenes and Croats are
of over 20% in Romania to the east. genetically distant from each other, even if they speak
related (South-Slavic) languages, while the Slovenes and
(4) Haplogroup E3b shows a south-north gradient and Hungarians are genetically close to each other, even if
its values are about 25% in Greece but only about 10% they speak unrelated languages. This complicated genet-
in the northernmost area of the Balkans. ic-linguistic relation may be explained by the fact that
there was previously a mostly homogeneous population
(5) A similar but weaker gradient concerns Haplogroup that spoke a common language, but then part of the
J2: its maximum value of about 20% in the south population shifted language and the original language
diminishes gradually to about 5% in the north. was replaced by Hungarian in some areas and by South-
Slavic in others.
On the basis of and the Balkan maps, the
following detailed genetic observations can be made: (2) The Slovenes, Croats, and Macedonians (all of
whom are Slavic-speaking) originate more strongly than
(1) The Slovenes belong to the Central European group others from the Ukrainian refuge. This is seen in their
with the Hungarians, their immediate geographic neigh- relatively high R1a values: 29.5% in the Slovenes,
bours. This is seen in the high R1b value, which is 31.8% in the Croats, and 35% in the Macedonians. In
The frequencies of Y Haplogroups R1a and R1b in the Balkans and some neighbouring
areas.
The frequencies of Y Haplogroups E3b and J2 in the Balkans and some neighbouring
areas.
the other Balkan populations the equivalent frequencies Slavic-speaking coastal areas that were perhaps the first
are considerably lower (5.8-16%). to receive farming in Europe. The lower E3b + J2 + G
values of the Macedonians (35%), Bulgarians (29%),
(3) High I-values (mostly I1b1-P37) are typical of the Bosnians (28%), Serbs (32%), Slovenes (17.0%), and
Croats (42%), Bosnians (48%), and Bulgars (42%). In Croats (11.1%) may indicate that the ancient areas of
the other Balkan populations, this value is lower (14.2- these populations were not equally suitable for early
30.4%). This can be interpreted as a possible indication farming.
of the fact that the Croats, Bosnians, and Bulgarians
originate from the Balkan refuge more often than their
neighbours.
In Bara et al. (2003) a detailed analysis of the male
(4) Strong indications of Early Farmers are seen in the populations of eleven Croatian localities is presented;
high E3b+J2+G values of the Turks (53.2%), Albanians see and for Haplogroups R1a,
(51.1%), and Greeks (48.3%). These areas are non- R1b, and I1b1-P37.
70
(3) The West Balkan peak area (frequencies over 50%)

of Haplogroup I1b1-P37 is in central Croatia near
Zagreb.
Accordingly, the Croatian men have come to their

present habitats from three directions, Slovenia and the
Iberian refuge, Hungary and the Ukrainian refuge, and
from the south; see .
The only present language of the Italian peninsula is

Italian, a Romance language, although prior to the time
of the Romans, there were others, such as Etruscan.
The frequencies of Y Haplogroup G in

the Balkans and some neighbouring areas.
Frequency zones of the frequencies of

Haplogroup R1a in Croatia.
Geographic locations of the eleven

Croatian localities in the study: six localities in the
mainland and five on the islands.
The maps allow the following generalizations about the

Croatian men:
(1) The R1a men arrived in Croatia mainly from the

areas now belonging to Hungary. Ultimately, these men
came from the Ukrainian refuge.
(2) The R1b men arrived in Croatia from the Iberian

refuge, mostly through Slovenia and an area now a part Frequency zones of the frequencies of
of Austria. Haplogroup R1b in Croatia.
Of the three large islands near the peninsula, Sicily and

Sardinia belong to Italy and are Italian-speaking. The
third island of the area, Corsica, belongs to France and The Wikipedia and Balanovsky maps show only one
is French-speaking. We first consider the Italian penin- strong gradient in the Italian peninsula: The frequencies
sula, and afterward, the three islands. of R1b are about 40% or more in the north and only
about 15-30% in the south. A candidate for a weak
gradient is offered by the “West-Balkan” Haplogroup
I1b1-P37. For this haplogroup, the frequencies are
1-5% in the south and practically non-existent (0-1%)
in the north.
More detailed information about the haplogroup fre-

quencies in various parts of the Italian peninsula is
found in . The tables are based on two
investigations, one by Scozzari et al. (2001) and the
other by Di Giacomo et al. (2003).
The seventeen geographic locations of are

shown in . The haplogroup frequencies of the
male populations of these localities are shown in
.
(1) As show in , the R1b-frequencies are higher

than 70% in northern Italy, but only about 25% in
southern Italy. In addition to this north-south gradient,
there are exceptionally low frequencies (11-18%) of
R1b in the southeast (Foggia and Brindisi). The frequen-
cy distribution of Haplogroup R1b from the study by
DiGiacomo (2003) is very similar, as shown in
The geographic locations of the seventeen

areas of
Frequencies of Y Haplogroups R1b and R1a in the Italian peninsula.

72
61.1 0 11.1 72.2 11.2 11.1 0 22.3 5.6 0 18

30.0 10.0 30.0 70.0 25.0 5.0 0 30.0 0 0 20
41.2 0 35.3 76.5 5.9 17.7 0 23.6 0 0 17
33.3 3.0 25.7 62.0 10.6 25.7 0 36.3 1.5 0 66
20.0 0 26.7 46.7 0 53.4 0 53.4 0 0 15
21.1 10.5 31.6 63.2 21.1 15.8 0 36.9 0 0 19
34.5 3.9 26.7 65.1 12.3 21.5 0 33.8 1.1 0 155
39.1 3.8 0 42.9 20.5 28.8 2.3? 51.6 5.4 0 131
19.6 1.6 55.6 76.8 7.5 12.5 0.3 20.3 0.9 4.1 331
73.3 6.7 80.0 10.0 10.0 10.0 30

45.5 9.1 4.5 59.1 9.1 27.2 36.3 4.5 22
76.2 4.8 81.0 2.4 9.5 4.8 16.7 2.4 42
48.3 6.9 55.2 24.1 10.3 10.3 44.7 29
25.7 5.7 8.6 40.0 11.4 34.3 5.7 51.4 8.6 35
45.0 45.0 15.0 30.0 45.0 10.0 20
41.4 6.9 10.3 58.6 3.4 20.6 6.9 30.9 10.3 29
26.1 2.2 8.7 37.0 17.4 26.1 10.9 54.4 8.7 46
29.2 2.1 6.3 37.6 12.5 27.2 14.6 54.3 8.3 28
11.1 18.5 29.6 11.1 44.4 14.8 70.3 27
27.6 3.4 3.4 34.4 24.1 37.8 61.9 3.4 29
30.0 10.0 5.0 45.0 20.0 25.0 10.0 55.0 20
18.4 5.3 13.2 36.9 26.3 23.7 7.9 57.9 5.3 38
40.0 4.0 44.0 36.0 12.0 4.0 52.0 4.0 25
33.3 12.5 45.8 25.0 12.5 4.2 41.7 12.5 24
25.9 3.7 7.4 37.0 11.1 44.4 55.5 7.4 27
24.2 24.2 27.3 36.4 3.0 66.7 9.1 53
36.4 3.4 6.5 44.3 15.8 25.2 6.3 47.3 6.3 524
27.5 3.9 3.9 5.9 31.4 11.8 49.1 11.8 3.9 51

23.1 0 37.2 10.3 5.1 14.1 29.5 5.1 5.2 78
50.0 0 8.8 14.7 2.9 11.8 29.4 2.9 8.8 34
33.5 1.3 16.6 10.3 13.1 12.6 36.0 6.6 6.0 153
Frequencies of Haplogroup R1b in the Frequencies of Haplogroup I in the

Italian peninsula. The map is a simplified version Italian peninsula.
of a map in Di Giacomo et al. (2003).
Frequencies of Haplogroup DE in the Frequencies of Haplogroup G2 in the

Italian peninsula. Most of Haplogroup DE is Italian peninsula.
represented by E3b in Italy.
74
Frequencies of Haplogroup J in the Italian Frequencies of Haplogroup Y* (”Other”)

peninsula. in the Italian peninsula, probably including men of
Haplogroups L and K2.
(2) For Haplogroup R1a, no equivalent gradient is

found. Instead, there seems to exist two separate fre-
quency peaks (both of the magnitude of about 5-10%);
one covers almost the entire northern half of Italy and The frequencies of seven haplogroups in Sicily, Sardinia,
the other the southeastern corner (Casarno and Bran- and Corsica are seen in . The following gener-
disi) of the peninsula. alization are possible on the basis of these data:
(3) shows that the distribution of Haplogroup I (1) Sicily has much higher percentages (5.9+31.4 =
is rather patchy. One general observation about the 37.3%) for the Early Farmers than the other two islands,
distribution of this haplogroup is that the frequencies whose equivalent percentages are only 10.3+5.1 =
are highest (about 20%) in Foggia and zero in the 15.4% and 14.7+2.9 = 17.6%. Another difference
extreme north (Valid Non and Garfognana) and the concerning the Early Farmers is that the ratio of the
extreme south (Reggio and Calabria). The Italian men frequencies of two of the haploggroups is different. The
of this haplogroup undoubtedly arrived from the Bal- “African” Haplogroup E3b is more frequent than the
kans, but the present uneven distribution seen today “Near Eastern” Haplogroup J2 in Sardinia and Corsica,
does not lend itself to simple explanations. but in Sicily J2 is much more frequent than E3b. This
difference may indicate a difference in the areas respon-
(4) The haplogroups of the Early Farmers are quite sible for the arrival of agriculture in the three islands.
common in Italy. The sum total of Haplogroups
DE+G2+J covers almost half (47%) of the Italian men of (2) Sicily has also higher F* frequencies (about 12%)
the peninsula. The areas of the highest frequencies are than the other two islands (about 3-5%). This may be
naturally in the south (as agriculture arrived from the another indication of relatively large number of men
east along the Mediterranean coast), but also the coastal arriving in Sicily by boat from the Middle East.
area close to France (Genoa) has quite high percentages
of these haplogroups. (3) A typical feature of Sardinia is the surprisingly high
frequency (almost 40%) of Haplogroup I (more pre-
cisely its subhaplogroup I1b1-P37). This may indicate

an early contact with the Western Balkans.
(4) About a half of the Corsican men represent the

“Iberian” Haplogroup R1b. In the other two islands,
the equivalent frequency is only about 25%. In this
respect, the Corsican men are typically West-Europeans
and many of them have, no doubt, come to the island
from France and Spain.
Iberia consists of three areas: Spain, Portugal, and the

Basque area. Linguistically, Iberia represents two lan-
Frequencies of Haplogroup R1b in the
Iberian peninsula.
The locations of the thirteen Iberian Frequencies of Haplogroup E in the

populations discussed in the text. Iberian peninsula.
60.0 11.3 71.3 7.0 11.0 18.0 4.5 5.2 155

59.1 9.2 68.3 13.5 13.7 27.2 4.5 0 22
53.5 14.3 57.8 3.6 17.9 21.5 10.7 0 28
55.8 14.7 70.5 11.1 14.7 25.8 3.7 0 27
42.5 0 42.5 26.8 15.4 42.2 0 15.3 26
53.3 5.5 58.8 15.5 12.0 27.1 1.8 11.0 109
61.5 3.4 65.9 10.0 8.7 18.7 3.3 15.1 60
63.1 0 63.1 31.6 5.3 36.9 0 0 19
58.4 5.7 64.1 10.0 5.8 15.8 4.3 15.8 70
54.8 12.9 67.3 16.2 9.7 25.9 3.2 3.2 31
52.4 33.3 85.7 4.8 9.5 14.3 0 0 21
89.0 8.8 97.8 2.2 0 2.2 0 0 45
75.1 6.2 81.3 0 ? 18.7 0 0 16
59.9 9.6 68.8 11.9 9.5 21.4 2.5 5.0 629
Note: R1b = R1 (xR1a), R1b-M153, R1b-SRY2627; I = I (xI1b2), I1b2; E = E*, E3a, E3b1,
E3b2, E3b3*, E3b3a; Others = BC*, DE*(xE3), F*, G, N3a, P*(xR1), R1a.
76
Frequencies of Haplogroup J in the Frequencies of Haplogroup I in the

Iberian peninsula. Iberian peninsula.
guage phyla, Indo-European and Basque. The Indo-Eu- (1) The average total of the frequencies of Haplogroup
ropean languages belong to the Romance group and R1b in Iberia is about 60%. The centre of this haplo-
represent two main languages, Spanish (Castilian) and group (89%) is in the Basque area but the frequencies
Portuguese. Spanish is often interpreted as containing are quite high (75%) also in Catalonia. In most parts of
two regional languages Galician and Catalan. the peninsula, the R1b-frequencies are about 50-60%.
The lowest frequencies (about 43%) are in the Malaga
The Y-chromosome haplogroups and their frequencies district in the southeastern corner of Iberia. There is a
are shown in . The frequency zones of the most north-south clinal gradient with higher values (over
common five haplogroups are seen in . The 80%) in the Northeast and lower values (close to 40%)
maps are based on . in the Southeast. The high values of Haplogroup R1b
reflect the Ice Age Iberian refuge, and the men of this
The subgroups contributing to the total R1b frequencies haplogroup can be considered the original inhabitants of
in are as follows: R1 (x R1a, R1b-M153, the peninsula (after the Neanderthals), the first of whom
R1b-SRY2627), (50.1%), R1b-M153 (2.9%), and R1b- arrived there about 35 kya.
SRT2627 (6.9%). Haplogroup R1a is not illustrated in
a separate map because it is so small in Iberia (frequency (2) The average of the frequencies of Haplogroup I in
in entire Iberia = 1.7%), but it is included as one of the Iberia is about 10%. The geographic distribution of this
haplogroups called ”other” in . haplogroup is peculiar in that the peak area (frequency
about 33%) is in the middle of the peninsula (Castile)
The overall frequencies of the subgroups of Haplogroup and the frequencies diminish as the distance from this
I in the entire Iberian area are as follows: I (x I-M26) area increases. In the next zone, frequencies vary from
(6.0%) and I-M26 (3.7%). In Castile, the frequency of about 12% to about 15%; in the zone still further away
both components of I are quite high, but the ”Sardinian” from Castile, the frequencies are 6-9%, and in the zone
Haplogroup I-M26 is exceptionally high (19%). furtherest away from Castile (in the northwestern and
southeastern corners of the peninsula) the frequencies
are very low (0-3 %).
The overall frequencies for all of Iberia for the compo-
nent subgroups of Haplogroup E are as follows: E3 (x (3) The average total of Haplogroup E in the peninsula
E3a, E3b-M78, E3b-M81, E3b-M34) (0.9%), E3a is about 12%. The haplogroup has two centres, one in
(0.6%), E3b-M78 (2.7%), E3b-M81 (5.5%), E3b- Galicia in the northwest (frequency about 32%) and the
M123 x E3b-M34 (0.1%), E3b-M34 (1.9%). other in Malaga in the southeast (frequency about 27%).
The men of clan E are ultimately of African origin, but
The frequencies in the entire Iberian for the components they have come to Iberia by two routes, some directly
of Haplogroup J are as follows: J x J2 (1.6%), J2 x from Africa across the Mediterranean, and others round
J2-M67 (5.6%), and J2-M67 (2.2%). the Mediterranean and through the Middle East. The
latter men were with the Early Farmers who brought
Several conclusions may be drawn from and agriculture to Iberia.
for Iberia:
. Frequencies of Y-chromosome haplogroup K

in the Iberian peninsula.
Haplogroups R1b and E, are, to some extent, comple-

mentary: The areas with high R1b values usually have
low E values and vice versa. This dichotomy results
from the population being originally R1b, while the E
people are more recent immigrants.
(4) The history of Haplogroup J is similar to that of

Haplogroup E in that it, too, belonged to the Early
Farmers arriving from the Middle East, and, according-
ly, Clan J, like Clan E, represents the newcomers to
Iberia (not the original inhabitants of the Iberian refuge). The area defined as ”Atlantic Europe” in
The average total of Haplogroup J is about 9% in Iberia, this article (darker shaded area).
61.0 0 13.6 74.6 6.7 10.1 3.4 20.2 0 3.4 Kivisild (1999)
50.0 5.0 25.0 80.0 11.0 5.0 5.0 21.0 0 0 0 Rosser (2000)
52.2 0 17.2 69.4 8.7 17.3 0 26.0 0 0 4.3 Semino (2000)
86.4 0 9.1 95.5 0 4.5 0 4.5 0 0 0 Semino (2000)
63.0 4.0 23.0 90.0 2.0 5.0 1.0 8.0 1 Rosser (2000)
60.0 4.8 25.7 90.5 2.4 6.2 0.8 9.4 Athey (2008)
57 3 22 82 6 9 2 17
67.6 11.8 17.6 97.0 2.9 2.9 99

66.7 5.1 10.2 82.0 5.1 2.6 5.2 12.9 5.1 125
30.8 15.4 11.4 57.6 19.2 15.4 34.4 7.7 109
55.0 10.8 13.1 78.9 9.1 6.0 1.7 16.7 1.7 2.6 333
78
and the frequencies form a south-north gradient. (5) Haplogroup K (x NO, P) is not very common in
has three zones: the frequencies of the southern zone Iberia as shown in . Its average frequency in the
are 14-18%, those of the middle zone 10-12% and those whole peninsula is about 3%, most of which is K2. The
of the northern zone 0-9%. The gradient suggests that maximum area is in the Cadiz area (west of Gibraltar)
agriculture spread in Iberia from the south to the north. which may mean that the men of this haplogroup first
arrived in southern Spain.
Frequencies of Y Haplogroup R1b in Frequencies of Y Haplogroup I in

Atlantic Europe. Atlantic Europe.
Frequencies of Y Haplogroup E3b in Frequencies of Y Haplogroup J2 in

Atlantic Europe. Atlantic Europe.
(1) The ”Iberian” Haplogroup R1b is very frequent in are summarized in . are based
Atlantic Europe. Its frequency is about 86% in the upon the table. The following generalizations about the
French Basque area and about 50-63% in the other British Isles data can be made:
continental areas of France and Belgium/Holland.
(1) The European maximum area of the ”Iberian” Hap-
(2) In the Atlantic Europe area, the ”Balkan” Haplo- logroup R1b is in Ireland. From there starts a west-east
group I is highest (about 27%) in Holland and slightly gradient that goes through Britain and continues on the
lower (14-25%) in Belgium and France; the I-frequency Continent. So, for example, frequencies are about 95%
is lowest (below 10%) among the Basques of France. in far northwest Ireland, 60% in the eastern parts of
There is a north-south gradient from Holland through England, about the same in Belgium and Holland, about
Belgium and France to the Basque area. 30-45% in Germany, and about 10-20% in Poland.
(3) The frequencies of Haplogroups E3b and J2 of the (2) The ”Ukrainian” Haplogroup R1a is relatively rare
Early Farmers are spread more or less uniformly across in the British Isles: its frequencies are usually below
the entire area under consideration. The average total 10%. There is, however, one exception: the frequencies
of E3b is about 6% and that of J2 about 8%. The only are about 10-20% in the northern islands. This can be
exception seems to be the very low (0-2%) frequency of seen as an indication of Scandinavian influence: in Nor-
E3b in the Basque area. way and Iceland, for example, the frequencies of R1a are
usually above 20%.
(3) In the central parts of eastern England, the frequen-

The data for Britain and Ireland, considered here as cies of Haplogroup I are higher (about 22-32%) than
”The British Isles,” are from Capelli, et al. (2003), and elsewhere in the British Isles (about 5-20%). This per-
66 23 10 99 0 83
64 19 15 98 0 2 121
80 6 14 100 0 51
66 9 25 100 0 99
79 5 13 97 0 2 44
80 2 10 92 7 7 41
86 4 7 97 0 2 42
73 3 18 94 4 4 2 96
68 8 18 94 3 2 5 1 90
70 13 16 99 2 2 62
57 4 32 93 4 4 2 46
64 5 28 87 6 6 12 70
71 2 18 91 4 4 8 1 84
66 4 19 89 5 2 7 4 57
89 1 4 94 4 1 5 1 80
86 4 11 101 0 76
90 0 9 99 0 43
60 4 32 95 3 2 5 121
91 2 4 97 3 3 59
65 8 22 95 4 4 2 51
76 4 11 91 4 5 9 55
74 1 18 93 1 5 6 1 80
73 4 14 91 4 4 8 1 73
79 8 12 99 2 2 52
66 3 24 93 4 2 6 2 128
70.9 5.8 16.2 95.1 1.9 2.0 3.9 0.8 0.1 0.0 1863
Note: Simplifications: JxJ2 + J2 = J; N3 removed (always zero); PxR removed (only
Orkney 2); KxPNO removed (only Llangefni 1); R1a1 = R1a; R1xR1a1 = R1b; FxIJK =G.
80
Frequencies of Y Haplogroup R1b in the Frequencies of Y Haplogroup R1a in the

British Isles. British Isles.
Frequencies of Y Haplogroup I in the Frequencies of Y Haplogroup E3b in the

British Isles. British Isles.
It is not certain where and when R1 split (via a mutation

in M173) from R. It was thought earlier that this split
took place in central Asia about 35 kya (Wells et al.
2001; Cordaux et al. 2004). According to a newer
interpretatiom, the mutation may have taken place in
India. This idea is supported by the fact that it is only in
India that the older variants of R, namely R*-M207 and
R2-M124 are commonly found. Another explanation
for India as the source area for R is the fact that India is
also a common initial area for P*, the ancestor of all R
variation (opinion of Richard Villems, private communi-
cation).
(2) Even before the rise of R1-M173, the

”grandfather” of R1, Clan F (defined by M-89) started
to expand in the Middle East, and about 21–28 kya Clan
I (defined by M170) started to spread outside the Middle
East. This clan was probably born in the Balkans (the
Balkan refuge) about 20 kya or earlier and it spread
from there to Central and Northern Europe during the
recolonization of Northern Europe. Clan I was divided
into several subclades, the main types of these being
”Scandinavian” I1a, ”Balkan” I1b1-P37, and
”German” I1b2-M223.
(3) At about the same time that Clan I was

founded, a Siberian Haplogroup N man gave rise to two
Frequencies of Y Haplogroup J in the subclans: ”Northeast European-Siberian” Clan N3 and
British Isles. ”Northwestern Siberian” Clan N2. The subgroups of N
are common in the northeastern corner of Europe as
well as throughout Siberia. They are practically nonex-
istent in Central, Western or Southern Europe.
centage is high (25%) also in the Hebrides. Most of the
instances of Haplogroup I belong to the ”Scandinavian” (4) Agriculture started to arrive in
Haplogroup I1a-M253. Europe about 10 kya. It arrived there in two possible
ways, either as demic diffusion or as cultural diffusion
(4) The frequencies of early farmers’ Haplogroups E3b or both. If agriculture was brought to Europe by Clans
and J are not very high (only about 2% for each) in the E3b, J2, and G, the men of these clans lived still outside
British Isles. Surprrisingly, the peak value of J (7%) is in Europe during the Ice Age; they did not live in any of the
Central Scotland. On the whole, there is a general European refuges, but in the Near East or its vicinity.
tendency according to which the frequencies of E3b+J According to the other possibility, the representatives of
are higher in the south and east and lower in the north these clans lived in the Balkan and/or Ukrainian refuge
and west. as early as the Ice Age, and represent the first Europeans
to learn how to cultivate soil and raise cattle.
In this review of the Y-chromosome evidence from all

parts of Europe, it is clear that European men came to When analysing the origins of European populations at
Europe in four main groups at different times: least three alternative approaches can be used: linguis-
tic, archaeological, and genetic.
(1) The first modern humans to arrive in
Europe (then occupied only by Neanderthals) were the (1) Earlier, linguists played a key role in studying the
descendants of an Asian man who was the founder of origin and early history of peoples. Using the present
Clan R1, defined by a mutation in the genetic marker and earlier regions of languages and the relationships
M173. The clan was later divided into two subclades, between the regions where they were spoken as their
”East European-Indian” R1a and ”West European” tools, linguists thought that they could arrive at the
R1b. original homes of languages or, in other words, they
thought they were able to answer the question
82
The special part played by lan- exerted an influence on the language situation in Europe.
guages was largely based on the concept of the nation In ancient times, there may have been many languages in
state, according to which nations were decided on the Europe, now extinct, about which we know nothing.
basis of the languages they spoke: the French were
primarily those who spoke French, the Estonians those Iberia
who spoke Estonian, etc. The in looking
for a common origin of peoples and languages, often The most plausible candidates for the ancient languages
even unquestioningly self-evident, was the idea that of the Iberian refuge are the Basque languages still
or "language determines the na- spoken by about half a million people in the Basque area
tion." However, the methods used by linguists have of Spain and France. Earlier, there were several lang-
their limitations, especially when it comes to time. The uages belonging to this language group, but mainly
farthest back in time that linguists can go is usually because of the intensive spread of IE languages in West-
regarded as 6,000-10,000 years. In other words, the ern Europe, the area of the Basque languages has shrunk
study of language takes us back no farther than Meso- ever since. It is probable that the entire Atlantic Coast
lithic time; the Palaeolithic era remains, as far as the was linguistically Basque during the Last Glacial Maxi-
roots of peoples are concerned, completely unstudied. mum (LGM) and the millennia after it. The area was
homogeneous also in respect to subsistence system and
(2) Later, especially in the 1970's and 1980's, archaeol- genetics: the men were reindeer hunters and their main
ogists have joined the numbers of those interested in the Y-chromosome haplogroup was R1b.
origin of peoples. Archaeologists are able to gain relia-
ble information from much earlier periods than 10,000 Siberia
years ago but they have been cautious in pronouncing
upon the origins of peoples. They often point to the fact It is a commonly accepted idea that the languages of the
that it is difficult to link archaeological cultures to “ancient mammoth hunters” of northeastern Europe
languages. They lived at a time when one had to know and northwestern Siberia were . It is possi-
what language a people spoke in order to be able to say ble that all these men occupied the entire northernmost
who the ancestors of the present population were. zone of Europe during the LGM and the period after it.
The populations had a common subsistence system and
(3) A decisive change came about when start- they were genetically homogeneous: they were mam-
ed in the 1980's to seriously study peoples' roots. Now moth hunters and their main Y-chromosome haplo-
came the time when people's origins were decided ac- group was N3.
cording to their genes rather than the language they
spoke. The geneticists, then, could construct two-di- The Ukraine
mensional trees for people, the one dimension of which
was the degree of relativity between peoples and the The men of the Ukrainian refuge, like those of the
other dimension time. These are in Siberian refuge, were mammoth hunters. They are com-
principle the same as the traditional of monly known for their houses made of mammoth bones.
the linguists; the difference is that instead of languages The language of these men may have the same
genes are used to identify peoples. language as that of the Siberian refuge; another
alternative is that it was . The IE branch
In this article, it has been my purpose to define popula- in question was the group consisting of
tions exclusively in genetic terms. To allow a compari- the GBS (Germanic, Baltic, and Slavic) languages. These
son of the old linguistic way of defining populations and language spread later (during the recolonization of
the new genetic way of defining them possible, I add Northern Europe) to the northern zone then occupied by
some concluding remarks about the assumptions con- the FU speakers from the Siberian refuge. The result was
cerning the languages spoken by ancient European pop- a rather strong FU substratum in all the GBS languages.
ulations. The time depth in many cases goes beyond the These men had arrived in Europe through the steppe
limits of linguistic facts, which means that the state- area between the Ural Mountains and the Caspian Sea;
ments in most cases are closer to assumptions than before that, they had occupied Central Asia and the
verified facts. Middle East/India. The main Y-chromosome haplo-
group of these men was R1a. Haplogroup R1a was
derived from the Middle East Haplogroup F through
mutations that gave rise to F > K > P > R > R1 > R1a.
Very little is known about the languages of the ancient
Europeans. Nevertheless, some hypotheses can be made The Balkans
about the languages. According to one view, each of the
four Ice Age refuges had its own language; in addition, The men of the Balkan refuge were more likely than
there were, of course, the languages of the southern those of any other to have spoken an early form of the
populations of the Middle East and Africa that may have Indo-European language. The IE language in question
would have given rise to the West-European group two linguistic types were assimilated but the two Y
consisting mainly of the Greek-Italic-Celtic languages. types, of course, remained separate.
One hypothesis is that IE languages were first brought
to Europe by the Early Farmers, displacing what had (5) A similar language shift may have taken place in
previously been all non-IE languages, but a more proba- northern Central Europe in the area of Proto-Germanic.
ble scenario is that IE came much earlier with the Hap- At least part of this area was earlier inhabited by men
logroup I men. In either case, the languages of the representing Haplogroup N3, and the language may
European Haplogroup I men shifted to the IE languages have been Finno-Ugric. The present Germanic languag-
of the Early Farmers during the Neolithic expansion out es (such as Slavic and Baltic) have a strong Finno-Ugric
of Anatolia. Genetically, the men of the Balkans repre- substratum (Wiik 2002).
sent Haplogroup I, which is a further development from
the Middle Eastern Haplogroup F. (6) Modern Hungarian men are genetically similar to
other Central Europeans with high R1b, R1a, and I1b
frequencies, but their language is Finno-Ugric. The
genetic-linguistic discrepancy can be solved by assuming
Contrary to the general way of thinking among tradi- language shifts in which the local Pannonian men
tional linguists, it is apparent that language shifts have accepted the Hungarian language of the newcomers as
been common during the time of modern man in Eu- their native language. The newcomers were the horse-
rope, and by comparing the genome and languages one riding hordes that came from the southern Ural Moun-
can make detailed assumptions about the language shifts tains to Pannonia in 500-895 AD. Hungarian men came
having taken place in Europe. At least the following from three different refuges: Iberia, resulting in high R1b
eleven major language shifts seem to have occurred in frequencies, the Balkans, resulting in significant I1b1-
Europe: P37 frequencies, and the Ukraine, resulting in high R1a
frequencies. Linguistically, all speak the same Hun-
(1) The men of the South-Slavic populations of the garian language and cannot be distinguished on this
Balkans are genetically from the Balkan refuge with high basis.
frequencies of Haplogroup I1b1-P37, but linguistically
they are from the Slavic group. In this case, the Balkan (7) Before the arrival of the Angles and Saxons, the
populations (whatever their original language) seem to language of most of those living in the British Isles was
have shifted their original language to a Slavic one. A Celtic. Today, Celtic languages are spoken only in the
strong indication of languages shifts is offered by the most remote areas of Ireland, Wales, and Scotland. A
existence of the ”Balkan Sprachbund” consisting of a wave of language shift Celtic > English has swept over
number of languages with unrelated vocabularies, but the British Isles during the last approximately sixteen
with similar grammatical and phonological features. hundred years.
(2) The emergence of the Romance languages is based on (8) A similar language shift that wiped off the Celtic
the language shift of the original local languages to language from Central Europe was more effective than
Latin. Depending on the original local language, the the one in the British Isles. According to the language
resulting language was, for example, French, Spanish, shift in question, a majority of the Central European
Portugese, Italian, or Romanian. Celts learned to speak a Germanic language and a mino-
rity learned to speak a West-Slavic (Polish, Czech, and
(3) In Central and Northern Russia, the original Finno- Slovak) language.
Ugric languages were replaced by Russian. The original
FU-speaking people learnt to speak Russian as their (9) In central and northern Finland, the speakers of the
native language. Saami language shifted their language to Finnish; only
the most northern Saami retained their original language
(4) An equivalent language shift took place in the Baltic and still today speak Saami.
area. The Latvian and Lithuanian men are genetically
partly from the Siberian refuge with a high N3 frequency (10) The Samoyeds of Northeastern Europe are geneti-
and partly from the Ukrainian refuge with high R1a. As cally different from all other Europeans, but their lan-
the languages of this area are today Baltic, a language guage is Uralic and related, for example, to Finnish. The
shift or, more precisely, a linguistic assimilation, must complicated genetic-linguistic situation is probably a
have taken place. The men who came originally from result of a language shift in which the Samoyeds came
the Siberian refuge must have shifted their Finno-Ugric into close contact with populations speaking a Finno-
language to a Baltic one: The Baltic area consisted earlier Ugric language. The result was a new language group,
of two genetic types (N3 and R1a) and two linguistic the Samoyedic languages, that are related to the Finno-
types (Finno-Ugric and Indo-European/Baltic); later the Ugric languages. Traditionally, the Finno-Ugric and
Samoyedic languages are regarded as ”Uralic.”
84
(11) In the Volga area, genetic and linguistic assimila- Francalacci P, Morelli L, Underhill PA, Lillie AS, Passarino G,
tion and mingling has been common in the Finno-Ugric Useli A, Madeddu R, Paoli G, Tofanelli S, Calo CM, Ghiani
and Turkic populations. The FU populations of the area ME, Varesi L, Memmi M, Vona G, Lin AA, Oefner P, Cavalli-
Sforza LL (2003) Peopling of three Mediterranean islands
are Mari, Mordvians, and Udmurtians and the Turkic
(Corsica, Sardinia, and Sicily) inferred by Y-chromosome
ones are Tatars and Chuvash, and to a lesser extent biallelic variability. .
Bashkirians.
Karlsson AO, Wallerström T, Götherström A, Holmlund G
(2006) Y-chromosome diversity in Sweden – A long-time
perspective. , 14:863-970.
http://home.comcast.net/~hapest5/index.html Kasperavièiûtë D, Kucinskas V, Stoneking M (2004) Y chro-

mosome and mitochondrial DNA variation in Lithuanians.
Haplogroup Predictor Program
Kayser M, Lao O, Anslinger K, Augustin C, Bargel G,

Edelmann J, Elias S, Heinrich M, Henke J, Henke L, Hohoff
C, Illing A, Jonkisz A, Kuzniar P, Lebioda A, Lessig R, Lewicki
Athey TW (2005) Haplogroup prediction using an allele-
frequency approach. , 1:1-7. Schmid D, Schmidt U, Schneider PM, Stradmann-
Bellinghausen B, Szibor R, Wegener R, Wozniak M,
Athey TW (2006) Haplogroup prediction from Y-STR values Zoledziewska M, Roewer L, Dobosz T, Ploski R (2005)
using a Bayesian-allele-frequency approach. Significant genetic differentiation between Poland and
, 2:34-39. Germany follows present-day political borders, as revealed by
Y-chromosome analysis. 117:428-443.
Athey TW (2008) Private communication. Dutch YHRD data
from five localities were pooled (n = 275) and analyzed using King R, Underhill PA (2002) Congruent distribution of Neo-
the Haplogroup Predictor program. lithic painted pottery and ceramic figurines with Y-chromo-
some lineages. 76:707-714.
Balanovsky O, Rootsi S, Pshenichnov A, Kivisild T, Chumosov
M, Evseeva I, Pocheshkhova E, Boldyreva M, Yankovsky N, Kivisild T et al (1999) An Indian Ancestry: a Key for Under-
Balanovska E, Villems R (2008) The sources of the Russian standing Human Diversity in Europe and Beyond.
patrilineal heritage in their Eurasian context. . McDonald
, 82:236-250. Institute Monographs, Cambridge, UK.
Bara L, Peri i M, Klari IM, Rootsi S, Jani ijevi B, Kivisild Laitinen V, Lahermo P, Sistonen P, Savontaus ML (2002)
T, Parik J, Rudan I, Villems R, Rudan P (2003) Y-chromo- Y-chromosomal diversity suggests that Baltic males share com-
somal heritage of Croatian population and its island isolates. mon Finno-Ugric-speaking forefathers.
. .
Capelli C, Redhead N, Abernethy JK, Gratrix F, Wilson JF, Lappalainen T, Koivumäki S, Salmela E, Huoponen K,
Moen T, Hervig T, Richards M, Stumpf MP, Underhill PA, Sistonen P, Savontaus ML, Lahermo P ( 2006) Regional
Bradshaw P, Shaha A, Thomas MG, Bradman N, Goldstein differences among the Finns: A Y-chromosomal perspective.
DB (2003) A Y chromosome census of the British Isles. 376:207-215.
, 13:979–984.
Marjanovic D, Fornarino S, Montagna S, Primorac D,
Cordaux R, Deepa E, Vishwanathan H, Stoneking M (2004) Hadziselimovic R, Vidovic S, Pojskic N, Battaglia V, Achilli A,
Genetic Evidence for the Demic Diffusion of Agriculture to Drobnic K, Andjelinovic S, Torroni A, Santachiara-Benerecetti
India. , 304:1125. AS, Semino O (2005) The Peopling of modern Bosnia-
Herzegovina: Y-chromosome haplogroups in the three main
DiGiacomo F, Luca F, Anagnou N, Ciavarella G, Corbo RM, ethnic groups. , 69:757-763.
Cresta M, Cucci F, DiStasi L, Agostiano V, Giparaki M,
Loutradis A, Mammi C, Michalodimitrakis EN, Papola F, Nasidze I, Ling EY, Quinque D, Dupanloup I, Cordaux R,
Pedicini G, Plata E, Terrenato L, Tofanelli S, Malaspina P, Rychkov S, Naumova O, Zhukova O, Sarraf-Zadegan N,
Novelletto A (2003) Clinal patterns of human Y chromosomal Naderi GA, Asgary S, Sardas S, Farhud DD, Sarkisian T,
diversity in continental Italy and Greece are dominated by drift Asadov C, Kerimov A, Stoneking M (2004) Mitochondrial
and founder effects. 28: 387-395. and Y-chromosome variation in the Caucasus.
68:205-221.
Dupuy BM, Stenersen M, Lu TT, Olaisen B (2005) Geograph-
ical heterogeneity of Y-chromosomal lineages in Norway.
164: 10-19. (2005) Review of Croatian genetic heritage as revealed by
mitochondrial DNA and Y chromosomal lineages.
Flores, C et al (2004) Reduced genetic structure of the Iberian 46:502-513.
peninsula revealed by Y-chromosome analysis: implications
for population demography. Eur J Hum Gen, 12, 855-863.
Rootsi, S et al (2002) The Roots of Peoples and Languages of Villems R, Kashyap VK (2006) A prehistory of Indian Y
Northern Eurasia IV. Oulu. chromosomes: Evaluating demic diffusion scenarios.
(USA), 103:843-848.
Rootsi S, Magri C, Kivisild T, Benuzzi G, Help H, Bermisheva
M, Kutuev I, Barac L, Pericic M, Balanovsky O, Pshenichnov Saukkonen P (2006)
A, Dion D, Grobei M, Zhivotovsky LA, Battaglia V, Achilli A, Yliopistopaino, Helsinki.
Al-Zahery N, Parik J, King R, Cinnioglu C, Khusnutdinova E,
Rudan P, Balanovska E, Scheffrahn W, Simonescu M, Brehm Scozzari R, Cruciani F, Pangrazio A, Santolamazza P, Vona G,
A, Goncalves R, Rosa A, Moisan JP, Chaventre A, Ferak V, Moral P, Latini V, Varesi L, Memmi MM, Romano V, De Leo
Furedi S, Oefner PJ, Shen P, Beckman L, Mikerezi I, Terzic R, G, Gennarelli M, Jaruzelska J, Villems R, Parik J, Macaulay V,
Primorac D, Cambon-Thomsen A, Krumina A, Torroni A, Torroni A (2001) Human Y-Chromosome Variation in the
Underhill PA, Santachiara-Benerecetti AS, Villems R, Semino Western Mediterranean Area: Implications for the Peopling of
O (2004) Phylogeography of Y-chromosome haplogroup I the Region. 62:871-884.
reveals distinct domains of prehistoric gene flow in Europe.
, 75:128-137. Semino O, Passarino G, Oefner PJ, Lin AA, Arbuzova S,
Beckman LE, De Benedictis G, Francalacci P, Kouvatsi A,
Rootsi S, Zhivotovsky LA, Baldovic M, Kayser M, Kutuev IA, Limborska S, Marcikiae M, Mika A, Mika B, Primorac D,
Khusainova R, Bermisheva MA, Gubina M, Fedorova SA, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Underhill PA
Ilumäe AM, Khusnutdinova EK, Voevoda MI, Osipova LP, (2000) The genetic legacy of paleolithic
Stoneking M, Lin AA, Ferak V, Parik J, Kivisild T, Underhill in extant Europeans: A Y chromosome perspective. ,
PA, Villems R (2006) A counter-clockwise northern route of .
the Y-chromosome haplogroup N from Southestern Asia to-
wards Europe. 15:204-11. Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds
CA, Chow CT, Lin AA, Mitra M, Sil SK, Ramesh A, Rani
Rosser ZH, Zerjal T, Hurles ME, Adojaan M, Alavantic D, MVU, Thakur CM, Cavalli-Sforza LL, Majumder PP, Under-
Amorim A, Amos W, Armenteros M, Arroyo E, Barbujani G, hill PA (2006) Polarity and Temporality of High-Resolution
Beckman G, Beckman L, Bertranpetit J, Bosch E, Bradley DG, Y-Chromosome Distributions in India Identify Both
Brede G, Cooper G, Côrte-Real HB, de Knijff P, Decorte R, Indigenous and Exogenous Expansions and Reveal Minor
Dubrova YE, Evgrafov O, Gilissen A, Glisic S, Gölge M, Hill Genetic Influence of Central Asian Pastoralists.
EW, Jeziorowska A, Kalaydjieva L, Kayser M, Kivisild T, , 78:202-221.
Kravchenko SA, Krumina A, Kucinskas V, Lavinha J, Livshits
LA, Malaspina P, Maria S, McElreavey K, Meitinger TA, Underhill PA, Kivisild T (2007) Use of Y Chromosome ans
Mikelsaar AV, Mitchell RJ, Nafa K, Nicholson J, Nørby S, Mitochondrial DNA population structure in Tracing Human
Pandya A, Parik J, Patsalis PC, Pereira L, Peterlin B, Pielberg migrations. , 41:539-564.
G, Prata MJ, Previderé C, Roewer L, Rootsi S, Rubinsztein
DC, Saillard J, Santos FR, Stefanescu G, Sykes BC, Tolun A, Wells RS, Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva
Villems R, Tyler-Smith C, Jobling MA (2000) Y-chromosomal I, Blue-Smith J, Jin L, Su B, Pitchappan R, Shanmugalakshmi
diversity in Europe is clinal and influenced primarily by geog- S, Balakrishnan K, Read M, Pearson NM, Zerjal T, Webster
raphy, rather than by language. 67:1526- MT, Zholoshvili I, Jamarjashvili E, Gambarov S, Nikbin B,
1543. Dostiev A, Aknazarov O, Zalloua P, Tsoy I, Kitaev M, Mirra-
khimov M, Chariev A, Bodmer WF (2001) The Eurasian
Sahoo S, Singh A, Himabindu G, Banerjee J, Sitalaximi T, heartland: a continental perspective on Y-chromosome diversi-
Gaikwad S, Trivedi R, Endicott P, Kivisild T, Metspalu M, ty. .

Where Did The European Men Come From

Uploaded by

Copyright:

Available Formats

Where Did The European Men Come From

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Where Did The European Men Come From

Uploaded by

Copyright:

Available Formats

Kalevi Wiik

on the development of the human family, specifically the

Nearly all European men can be classified into one of the

A phylogenetic tree of about forty Y-chromosome haplogroups and subhaplogroups in

smaller circles represent locations of the peak areas of

Population Old Europeans Early Farmers Other

agriculture expanded as the indigenous people acquired

The geographic distribution of Haplogroup R1b. Source:

The total geographic distribution of Haplogroup R1a. Source:

The main areas of concentration of The eastern extentions of Haplogroup R1a

The geographic distribution of all the subhaplogroups of N.

Frequencies of Haplogroup N3 in Eurasia.

Frequencies of Haplogroup N2 in Euraasia.

The development of Haplogroup N from F through K and

The geographical distribution of Haplogroup E (including all of its subhaplogroups).

Clan N3 extended to the west and east; today, Clan N2

(5) From the European perspective, Haplogroups E3b,

(a) Haplogroup E3b is a branch of the ”African” Haplo-

(b) The source area of the ”Near Eastern” Haplogroup

(c) The source area of Haplogroup G was perhaps in

Haplogroup E originated in Africa. This is true also of

The geographic distribution of Haplogroup J2. Source: Sengupta et al. (2006).

of some other parts of the Balkans and southern Italy.

It is likely that there is at least some interdependence

The Indo-European languages represent seven groups:

The question of which language a population speaks

The approximate areas of European language groups:

model used than by genetic reality. The maps represent-

Frequencies of the Iberian Haplogroup R1b in Europe.

Frequencies of the Ukrainian Haplogroup R1a.

Frequencies of the Scandinavian Haplogroup I1a, a aubhaplogroup of the

Frequencies of the German Haplogroup I1b2-M223,

Frequencies of the West Balkan Haplogroup I1b1-P37, subhaplogroup of

Frequencies of the Northeast European Haplogroup N3, subhaplo-

Frequencies of the Samoyed Haplogroup N2,

Frequencies of the Early Farmers Haplogroup E3b, subhaplogroup of

Frequencies of the Early Farmers Haplogroup G in Europe.

The total area of Central Europe is shown in

is a compilation of haplogroup frequencies for

bear witness to the fact that agriculture

R1b R1a1 I N3 OE E3b DE J2 G EF F P K Other n

1 Tyrol 59.4 12.5 6.2 78.1 15.6 6.3 21.9

1 Lausanne 68.8 9.3 12.5 90.6 6.3 3.1 9.4

The approximate locations of the Central

Frequencies of Haplogroups R1b and R1a in Central Europe.

Frequency of Haplogroup I in Central Europe.

Frequencies of Haplogroups E3b and J2 in Central Europe.

shows the most frequent haplogroup at various

(3) The frequencies of the Balkan Haplogroup I in

In addition, all three main types of the male populations

As seen in the maps, the three genetic types represent

1 43.7 22.4 33.1 0.3 99.5 0 0 0 0 0 0

5 6.8 14.2 33.8 42.6 97.4 0 0 0 0 0 0

17.5 12.5 45.0 5.0 10.0 90.0 1.8 0 0 0 0 5.0 5.0 40

26.8 27.1 34.7 10.6 0.8

0 8.3 16.7 2.0 70.8 0 2.1 48

3.4 19.0 46.6 0 25.9 3.4 1.7 58

6.2 8.3 52.1 2.0 27.1 0 4.2 48

13.0 20.4 46.3 0 0 79.7 7.4 5.7 5.6 18.7