J Hum Genet (2007) 52:406414

DOI 10.1007/s10038-007-0125-6

ORIGINAL ARTICLE

Y-STR variation among Slavs: evidence for the Slavic

homeland in the middle Dnieper basin
Krzysztof Rebaa Alexei I. Mikulich Iosif S. Tsybovsky
Daniela Sivakova Zuzana Dzupinkova
Aneta Szczerkowska-Dobosz Zofia Szczerkowska

Received: 15 September 2006 / Accepted: 7 February 2007 / Published online: 16 March 2007

The Japan Society of Human Genetics and Springer 2007

Abstract A set of 18 Y-chromosomal microsatellite loci

was analysed in 568 males from Poland, Slovakia and three
regions of Belarus. The results were compared to data
available for 2,937 Y chromosome samples from 20 other
Slavic populations. Lack of relationship between linguistic,
geographic and historical relations between Slavic populations and Y-short tandem repeat (STR) haplotype distribution was observed. Two genetically distant groups of
Slavic populations were revealed: one encompassing all
Western-Slavic, Eastern-Slavic, and two Southern-Slavic
populations, and one encompassing all remaining Southern

K. Rebaa (&)
Z. Szczerkowska

Department of Forensic Medicine,
Medical University of Gdansk,
ul. Marii Skodowskiej-Curie 3A,
80-210 Gdansk, Poland
e-mail: [email protected]
A. I. Mikulich
Institute for the Study of Arts, Ethnography and Folklore,
National Academy of Sciences, Minsk, Belarus
I. S. Tsybovsky
Institute of Problems of Criminology,
Criminalistics and Forensic Expertise, Minsk, Belarus
D. Sivakova
Department of Anthropology, Comenius University,
Bratislava, Slovakia
Z. Dzupinkova
Department of Experimental and Applied Genetics,
Institute of Preventive and Clinical Medicine,
Slovak Medical University, Bratislava, Slovakia
A. Szczerkowska-Dobosz
Department of Dermatology,
Medical University, Gdansk, Poland

123

Slavs. An analysis of molecular variance (AMOVA) based

on Y-chromosomal STRs showed that the variation observed between the two population groups was 4.3%, and
was higher than the level of genetic variance among populations within the groups (1.2%). Homogeneity of northern Slavic paternal lineages in Europe was shown to stretch
from the Alps to the upper Volga and involve ethnicities
speaking completely different branches of Slavic languages. The central position of the population of Ukraine
in the network of insignificant AMOVA comparisons, and
the lack of traces of significant contribution of ancient
tribes inhabiting present-day Poland to the gene pool of
Eastern and Southern Slavs, support hypothesis placing the
earliest known homeland of Slavs in the middle Dnieper
basin.
Keywords Y chromosome
Microsatellites

Slavic populations

Introduction
Since the human Y chromosome is characterised by the
presence of the largest non-recombining region in the
whole human genome, sensitivity of genetic variation to
drift phenomena, a unique inheritance pattern and specificity to males, its polymorphism has been widely studied
by researchers interested in human evolution and forensic
geneticists (Jobling and Tyler-Smith 2003; Butler 2003).
Depending on the time scales of the population history
events, different types of polymorphic markers abundant
on the Y chromosome are available for research. Analysis
of slowly evolving Y-chromosomal biallelic polymorphisms have enabled deeper insight into prehistoric population movements and colonisation waves in Europe

J Hum Genet (2007) 52:406414

(Semino et al. 2000; Rootsi et al. 2004). On the other hand,

Y-chromosomal short tandem repeats (Y-STRs) are characterised by a relatively high mutation rate and seem to be
much more suitable for genetic studies of more-recent
events. Although the combination between Y-chromosomal microsatellites and biallelic polymorphisms yields
the highest level of resolution and a means of clarifying
complex genetic histories (Weale et al. 2001, 2002; Das
et al. 2004), Y-STR data alone also provide very useful
information for analyses of interpopulation diversity and
have been widely applied in resolving differentiation of
various human populations (Kayser et al. 2001; Ploski et al.
2002; Caglia` et al. 2003; Roewer et al. 2005; Immel et al.
2006).
The Indo-European linguistic family has the largest
number of speakers of the recognised families of languages
in the world today. In this family, Slavic languages form a
group of closely related languages with close to 250 million speakers worldwide (Schenker 1995). In Europe, Slavs
are the most numerous ethnic and linguistic group of
peoples, residing chiefly in eastern and southeastern Europe but also extending across northern Asia to the Pacific
Ocean. The early medieval great migrations in Europe utterly changed the ethnic and linguistic situation of the
continent and spread the Slavic settlement in the fifth to
sixth centuries over the major part of Eastern Europe,
leading to the ethno-cultural subdivision of the primarily
united Proto-Slavic community (Encyclopdia Britannica
2006). Nowadays, from the linguistic, cultural and geographic point of view, Slavs are customarily divided into
three major subgroups: Eastern Slavs (Belarusians, Russians, Ukrainians), Western Slavs (Poles, Slovaks, Czechs,
Lusatians), and Southern Slavs (Slovenes, Croats, Bosnians, Montenegrins, Serbs, Macedonians, Bulgarians).
Since the ethno-cultural subdivision of Slavs and the
formation of modern Slavic nations took place quite recently, and significant differences in Y-STR haplotype
distribution exist even between closely related human
populations (Roewer et al. 1996), hypervariable Y-chromosomal microsatellites seem to be markers of choice for
the study of mutual relations between different Slavic
ethnic groups. So far, it was demonstrated that Slavic YSTR haplotype paternal lineages form a separate branch
in a phylogenetic tree of European populations (Roewer
et al. 2001, 2005). However, no comprehensive analysis
of interpopulation Y-STR haplotype variation between
different Slavic groups was available. This study has
focused on providing a phylogenetic overview of closely
related Central-Eastern European populations of Poland,
Slovakia and Belarus to analyse their relationship with
each other and with other Slavic populations, and to
investigate how these relations reflect Slavs historical
migrations.

407

Materials and methods

Eighteen Y-chromosomal microsatellite loci: DYS19,
DYS388, DYS389I, DYS389II, DYS390, DYS391,
DYS392, DYS393, DYS426, DYS437, DYS438, DYS439,
DYS460, GATA H4.1, DYS385 a/b, and YCAII a/b, were
genotyped in randomly selected, unrelated Poles (n = 208),
Slovaks (n = 164) and Belarusians (n = 196) by means of a
multiplex polymerase chain reaction (PCR) and capillary
electrophoresis, as previously described (Rebaa and
Szczerkowska 2005). The Belarusian population included
samples from three distinct regions: 86 males from southern Belarus, 57 males from central Belarus, and 53 males
from northern Belarus. Additionally, all Belarusian samples were genotyped at the M46 (Tat) locus by a PCR
restriction fragment-length polymorphism (RFLP) method
(Kayser et al. 2005). The products of amplification of the
M46 locus were digested with Hsp92II restriction endonuclease (Rybakowski et al. 2002), separated by polyacrylamide gel electrophoresis, and visualised by silver
staining. The Y-STR haplotype data for other 20 Slavic
(n = 2,937) and nine neighbouring non-Slavic (n = 1,428)
populations were obtained either from the Y chromosome
haplotype reference database (http://www.ystr.org) (Roewer et al. 2001) or from the literature (Pepinski et al.
2004a, b; Lauc et al. 2005; Lovrecic et al. 2005; Marjanovic et al. 2005; Pericic et al. 2005; Spiroski et al. 2005;
Lessig et al. 2006; Rodig et al. 2007).
Analysis of molecular variance (AMOVA) was performed with the use of Arlequin 3.1 software (Excoffier
et al. 2005) to calculate matrices of pairwise FST and RST
values between populations. Associated probability values
were estimated from 10,100 permutations. Linearised FST
and RST values (Slatkin 1995) were applied to build a
neighbour-joining tree using the options NEIGHBOUR and
DRAWTREE in the PHYLIP package (Felsenstein 2004),
and to conduct a multidimensional scaling analysis using
the STATISTICA 7.1 software (StatSoft). In all calculations, DYS389 was considered as a haplotype of two
independent loci: DYS389I (repetitive stretches: p + q)
and DYS389II-I (repetitive stretches: m + n) (Rolf et al.
1998), while microvariants, null alleles and locus multiplications were treated as missing data.

Results and discussion

AMOVA of 18-locus Y chromosome haplotypes revealed
significant differences between the populations of Belarus,
Poland and Slovakia (FST = 0.0100; P < 0.0001) as well as
between the three Belarusian subpopulations (FST =
0.0084; P = 0.0034). Analysis of pairwise FST values for
18 Y-STRs demonstrated that heterogeneity within the

123

408

Belarusian population was caused by differences between

northern and central Belarus while the remaining pairwise
comparisons did not yield statistically significant FST values. The most outstanding populations were those of Poland and northern Belarus, while populations of central
Belarus, southern Belarus and Slovakia were genetically
indistinguishable. The Polish population was shown to be
distinct from all studied populations (P = 0.01 for comparison with Slovakia, P < 0.005 for all other comparisons), while in the case of northern Belarus, the only
insignificant FST value was for a comparison with the
southern part of the country.
Our results have demonstrated that the Y-STR haplotype
distribution does not reflect the linguistic and/or historical
affiliations between Slavic populations. The Polish paternal
lineages revealed by Y-chromosomal microsatellite haplotype analysis were previously reported to be distant from
a number of non-Slavic European populations and Slavicspeaking Muscovites (Ploski et al. 2002), but no Slavic
populations inhabiting Polands geographic neighbours
were taken into consideration. Despite the usage of very
closely related languages by Poles and Slovaks as well as
over half a millennium of a common state bordering the
majority of Poles and Belarusians, the Polish Y-STR haplotype heritage was shown to be distinct from that of both
neighbouring Slavic nations. On the other hand, analysis of
Y chromosome haplotypes defined by 18 loci has revealed
genetic homogeneity between Slovaks and two subpopulations of Belarus, although both populations are geographically distant and isolated, speak tongues belonging to
separate branches of the Slavic language group, and have
never shared common state borders throughout their histories.
Since for the majority of other Slavic populations, only
nine-locus haplotypes (DYS19, DYS389I, DYS389II-I,
DYS390, DYS391, DYS392, DYS393, and DYS385 a/b)
were available, only these loci were used for further
analysis. A comparison of our Polish haplotypes defined by
the selected nine loci with data for six Polish regional
subpopulations from the Y chromosome haplotype reference database confirmed previously observed homogeneity
of Polish subpopulations (FST = 0.0003; P = 0.66) (Ploski et al. 2002), and in AMOVA, all Polish samples were
combined. Results revealed four clusters of Slavic populations connected by a network of statistically insignificant
FST values (P > 0.05): (1) all Western-Slavic and EasternSlavic populations, Slovenes and western Croats; (2) Lusatians; (3) Southern-Slavic northern Croats and Bosnians;
(4) Southern-Slavic Serbs, Macedonians, and Bulgarians
(Table 1). However, at the significance level of 0.01, only
one such cluster involving all Slavic populations was disclosed. The distinctiveness of Southern-Slavic populations
was observed as a separate branch in a neighbour-joining

123

J Hum Genet (2007) 52:406414

tree, while multidimensional scaling has displayed a nucleus of seven genetically indistinguishable populations
with very small relative genetic distances, which involved
population samples of Eastern-Slavic (Ukrainian, Russian,
Belarusian), Western-Slavic (Slovak) as well as SouthernSlavic (Slovene) origin (Fig. 1).
Since there is no clear consensus over the accuracy of
different statistical parameters estimating genetic distances
between populations in studies using microsatellite markers, both a classical allele frequency-based differentiation
estimator (FST) and its stepwise mutation model-based
analogue (RST) are commonly reported (Balloux and Lugon-Moulin 2002). Therefore, we applied both distance
methods to assess genetic relations between various Slavic
and non-Slavic populations. In accordance with results
obtained for autosomal STRs in various human populations
(Perez-Lezaun et al. 1997; Destro-Bisol et al. 2000) and for
Y-chromosomal microsatellites in sub-Saharan Africans
(Caglia` et al. 2003), the pattern of Y-STR interpopulation
diversity among Slavs and neighbouring populations, based
on FST values, appeared to be congruent with known ethnohistorical relationships, while that based on RST values
revealed unexpected and unconvincing population affinities.
Both the multidimensional scaling plot and the neighbour-joining tree, based on the FST values, revealed genetic
proximity between related populations: (1) Germans from
Bavaria and Saxony, (2) Italians from Latium and Veneto,
(3) Turks from Anatolia and Bulgaria, and (4) Balts from
Latvia and Lithuania (Fig. 1). The FST-based results were
consistent with expectations also in case of three isolated
Slavic populations: (1) Lusatians from southeastern Germany, who are descendants of Slavic tribes that have
inhabited the lands between the Elbe and Oder rivers since
the fifth century (Encyclopdia Britannica 2006), (2)
Polish Belarusians, who colonised parts of Podlachia
(northeastern Poland) in the 15th to 16th centuries after
arrival from the Hrodna region (Wisniewski 1964), and (3)
a community of Russian settlers (Old Believers), who arrived in Podlachia in the eighteenth century from the Pskov
and Novgorod regions (Grek-Pabisowa 1968). All three
ethnic groups were shown to be homogeneous with only
one of all compared populations (Table 1), representing
their population of origin. Lusatians revealed Y-STR
homogeneity with the neighbouring population inhabiting
areas from which they are supposed to migrate, i.e. with
Poles. In the case of Podlachian Belarusians, such a population of origin was geographically the closest population
of central Belarus (involving the city of Hrodna), while in
the case of Polish Old Believers it was Russians from the
Novgorod region.
On the other hand, in the RST-based multidimensional
scaling plot and the neighbour-joining tree, only the two

0.0062 0.0215

0.0040 0.0068

0.0962 0.0326

0.0543 0.0188

BeN

BeC

0.0006 0.0132

RuMos 0.0479 0.0084

0.0946 0.0681

0.0642 0.0541

0.0825 0.0638

0.0935 0.0738

Ma

Bu

0.0336 0.0166
0.0583 0.0357

CroW
CroN

Se

0.0379 0.0073

Bo

0.0392 0.0042

Ukr

Slvn

0.0501 0.0144

0.0087 0.0036

RuNov 0.0562 0.0134

RuVla

0.0216 0.0188

0.0623 0.0232

RuPdl

0.0003

0.0017

0.0127

0.0230

0.0075

0.0037 0.0038

0.0509 0.0879

0.0430 0.0751

0.0351 0.0744

0.0434 0.0680

0.0053 0.0424
0.0154 0.0413

0.0016 0.0245

0.0422

0.0420

0.0326

0.0546

0.0091
0.0219

0.0024

0.0508 0.0542

0.0420 0.0530

0.0353 0.0441

0.0297 0.0595

0.0115 0.0158
0.0119 0.0316

0.0078 0.0188

0.0054 0.0024 0.0164

0.0066 0.0132

0.0003 0.0157

0.0034 0.0243

0.0210 0.0348

0.0171

0.0003

0.1404 0.0564

0.1243 0.0001

0.1174 0.0000

0.0000 0.0000

0.1132

0.1077

0.0956

0.1051

0.0516
0.0703

0.0271

0.0143

0.0247

0.0201

0.0068

0.0000

0.0006

0.0012

0.0048

0.0000

0.0000

0.0000

0.7386

0.1632 0.1102

0.0002 0.0000

0.0009 0.0004

0.3949 0.0431

0.3186 0.4992

0.0372 0.0126

0.5154 0.0168

0.0050 0.0000

0.0001 0.0001

0.0803

0.0657

0.0610

0.0646

0.0258
0.0378

0.0430 0.0516

0.0401 0.0447

0.0328 0.0392

0.0435 0.0526

0.0050 0.0109
0.0171 0.0232

0.0004 0.0069

0.0023 0.0012 0.0045

0.0120

0.5272

0.0605 0.0537

0.0509 0.0468

0.0439 0.0397

0.0483 0.0525

0.0098 0.0035
0.0193 0.0178

0.0009

0.0960 0.0181

0.5456 0.3909

0.2940 0.0279

0.0039 0.0000

0.0012 0.0000

0.6576 0.0352

0.1489 0.2622

0.2224 0.0006

0.8820 0.6930

0.0557 0.0013

0.0220

0.0191

0.0132

0.0268

0.0064

0.1419

0.0246

0.0045

0.1102

0.0009

0.0000

0.0002

0.0138

0.0558

0.0000

0.0544

0.0001

0.0027

0.0205

0.0083

0.0075

0.0044

0.0396

0.0000

0.0008

0.0000

0.0009

0.0000

0.0000

0.0000

0.0062

0.0013

0.0000

0.0000

0.0000

0.0000

0.0197

0.0074

0.0103

0.0001
0.0975

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0001

0.0000

0.0000

0.0000

0.0000

0.0000

0.4738

0.0161 0.0208

0.0023 0.0031

0.0003

0.0032 0.0000
0.0207 0.0034

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0000 0.0000

0.0891

0.2041

0.0000

0.0001
0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

0.0000

Slvn
CroW
CroN
Bo
Se
Ma
Bu
n = 121 n = 101 n = 150 n = 100 n = 114 n = 234 n = 122

0.0023 0.0013

RuVla Ukr
n = 152 n = 82

0.0061 0.0023

0.0055

0.1002

0.0007

0.2481

0.0554

0.2636

0.0530

0.0055

0.0003

BePdl
RuPdl RuNov RuMos
n = 157 n = 127 n = 50 n = 85

0.0000 0.0001

BeS
n = 86

0.0059

0.0129

0.0884

0.0003

0.0064 0.0140 0.0008

0.0144 0.0300

0.0558 0.0181

0.0453 0.0266

BeS

0.0185

0.0026

BePdl

0.0280 0.0032

Slvk

0.0036 0.0000

0.0186 0.0000

Po

0.0336

0.0139

Lu

Lu
Po
Slvk
BeN
BeC
n = 29 n = 1521 n = 164 n = 53 n = 57

Table 1 Y-STR haplotype pairwise FST values below the diagonal and corresponding P values above the diagonal (P values > 0.05 are bold) for 19 Slavic populations

J Hum Genet (2007) 52:406414

409

123

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J Hum Genet (2007) 52:406414

Fig. 1 Two-dimensional plot

obtained from multidimensional
scaling, and a neighbour-joining
tree, based on pairwise FST
values for nine-locus Y-STR
haplotypes observed in 19
Slavic and 9 non-Slavic
populations. Ellipses are traced
around genetically
homogeneous northern Slavic
populations and clusters of nonSlavic populations with known
ethno-historical affiliations.
Arrows indicate historically
proved directions of migrations.
A dotted line connects
populations with a disputed
direction of migration, which
inhabit areas designated
according to the various sources
as the Slavic homeland

Baltic populations were clearly isolated from unrelated

ethnicities, while separation of German, Italian and Turkish
populations was no longer visible (Fig. 2). Moreover, statistically insignificant P values for RST genetic distances
were observed between historically, linguistically, and
geographically unrelated populations such as Turks and
Italians from Latium, or Belarusians and Germans from
Saxony. In the case of the three isolated Slavic populations,
genetic homogeneity restricted only to the populations of
origin, as observed for FST values, was lost in the RST
method. Thus, the FST genetic distances reflect interpopulation relationships between the compared populations
much better than their stepwise-based analogues. This
indicates that the genetic differences in the Y-STR haplotype distribution between the Slavic populations and their
nearest neighbours are caused mainly by drift, and is explained by the fact that a period of time since the differentiation of the compared populations was too short to
allow for an effective impact of mutations on Y-chromosome variation (Perez-Lezaun et al. 1997; Caglia` et al.

123

2003). Therefore, in further analysis, we focused only on

the results obtained by the classical allele frequency-based
approach for estimation of Y-STR genetic distances.
Comprehensive analysis of Slavic Y-chromosomal microsatellite haplotypes on a European scale confirmed
previous observations for 18 Y-STR loci in the Polish,
Slovak and Belarusian populations: no relationship between the customary linguistic division of Slavs and the YSTR haplotype distribution was disclosed. The most
apparent genetic distance was found between the northern
(Eastern and Western) and Southern Slavs, who at the end
of the 9th century were separated by the invasion of FinnoUgric Hungarians. The AMOVA showed that the variation
observed between both population groups was 4.3%
(FCT = 0.0428; P = 0.0003), which was higher than the
level of genetic variance among populations within the
groups (1.2%; FSC = 0.0130; P < 0.0001). This difference
was even more profound when the RST-based distance
method was applied: genetic variation between both population clusters was 19.8% (RCT = 0.1984; P = 0.0002)

J Hum Genet (2007) 52:406414

411

Fig. 2 Two-dimensional plot

obtained from
multidimensional scaling, and a
neighbour-joining tree, based on
pairwise RST values for ninelocus Y-STR haplotypes
observed in 19 Slavic and nine
non-Slavic populations. An
ellipse is traced around a cluster
of populations with known
ethno-historical affiliations
Lu Lusatians; Po Poles; Slvk
Slovaks; BeN, BeC, BeS, BePdl
Belarusians from northern
Belarus, central Belarus,
southern Belarus, Podlachia;
RuPdl, RuNov, RuMos, RuVla
Russians from Podlachia (Old
Believers), Novgorod region,
Moscow region, Vladivostok
region; Ukr Ukrainians; Slvn
Slovenes; CroW, CroN Croats
from western Croatia, northern
Croatia (Zagreb region);
Bo Bosnians; Se Serbs; Ma
Macedonians; Bu Bulgarians;
Lit Lithuanians; Lat Latvians;
GeBav, GeSax Germans from
Bavaria, Saxony (Dresden
region); ItLat, ItVen Italians
from Latium, Veneto;
Hun Hungarians; TuAn, TuBu
Turks from Anatolia,
Bulgaria

while the interpopulation variance within the groups was

only 1.8% (RSC = 0.0228; P < 0.0001). The observed
northern Slavic Y-STR genetic homogeneity extends from
Slovakia and Ukraine to parts of Russia and Belarus, but
also involves Southern-Slavic populations of Slovenia and
western Croatia, and is the most probably due to a homogeneous genetic substrate inherited from the ancestral
Slavic population. However, due to the Y-STR proximity
of linguistically and geographically Southern-Slavic Slovenes and western Croats to the northern Slavic branch, the
observed genetic differentiation cannot simply be explained by the separation of both Slavic-speaking groups
by the non-Slavic Romanians, Hungarians, and Germanspeaking Austrians. A similar difference has been previously reported between Bulgarians and a few other Slavic
populations (Roewer et al. 2005), and our results demonstrate that other Southern-Slavic populations, namely
Macedonians, Serbs, Bosnians, and northern Croats are
genetically distinct from their northern linguistic relatives

as well. Roewer et al. (2005) attributed a possible explanation for these differences to the admixture of Y chromosomes of Finno-Ugric and Turkic-speaking peoples who
had invaded and settled in the Danube basin and the Balkans. However, we found that the only population that
revealed an insignificant FST value in comparison with the
Finno-Ugric Hungarians was the population of western
Croatia, and this putative admixture did not significantly
affect the Y-STR proximity of western Croats to Eastern
and Western Slavs (Table 1). All other FST values obtained
for comparison of nine-locus haplotypes of Hungarians and
Turks from Bulgaria and Anatolia, with those of 19 Slavic
populations appeared to be statistically significant
(P = 0.01 for a comparison of Hungarians with the population of central Belarus, P = 0.002 for a comparison of
Bulgarian Turks with Bulgarians, P 0.0003 for all other
comparisons). Thus, the contribution of the Y chromosomes of peoples who settled in the region before the
Slavic expansion to the genetic heritage of Southern Slavs

123

412

is the most likely explanation for this phenomenon. On the

other hand, our results indicate no significant genetic traces
of pre-sixth-century inhabitants of present-day Slovenia in
the Slovene Y chromosome genetic pool.
Although the existence of the Balto-Slavic linguistic
community, or at least territorial contiguity of Proto-Baltic
and Proto-Slavic in the past, is generally accepted
(Schenker 1995), AMOVA revealed significant differences
in Y-STR distribution between Slavic and Baltic populations (P < 0.005 for all pairwise comparisons), which is
likely to result from the previously observed different Ychromosomal haplogroup distribution (Rosser et al. 2000).
The Baltic populations are characterised by the high
incidence of the Y-chromosomal haplogroup N3 (47%
among Lithuanians, 32% among Latvians) (Rosser et al.
2000; Zerjal et al. 2001). Its distribution pattern in Slavic
populations indicates that Proto-Slavs did not carry this
lineage at a substantial frequency, since it is relatively rare
among Slavs and at high frequency was observed only in
some Russian subpopulations (Malyarchuk et al. 2004).
Since at the significance level equal to 0.01, the only
population that did not yield statistically significant FST
values for comparisons with other Slavic and both Baltic
populations was the Slavic-speaking population of northern Belarus, we estimated haplogroup N3 frequencies in
the three Belarusian subpopulations. The results suggest
that the uniqueness of the northern Belarusian population
is most likely due to the high incidence of Y chromosomes from the haplogroup N3 (18.9%), which has half
the frequency in central and southern Belarus (8.8 and
8.1%, respectively). Therefore, although the early ethnogenesis of the Belarusian nation has customarily been
linked to the gradual Slavicisation of the homogeneous
Baltic substrate on the territory of present-day Belarus
(Sedov 1970), only northern Belarus seems to be a transient area for the Baltic and Slavic settlement. Apart from
Balts, the N3 Y chromosomes are also prevalent among
Finno-Ugrians (Rosser et al. 2000; Zerjal et al. 2001), but
it was found that Y-STR haplotypes from the haplogroup
N3 differ in Baltic and Finno-Ugric populations, most
likely due to two distinct migration waves of people
carrying N3 Y chromosomes (Zerjal et al. 2001; Kasperavici
ute et al. 2004). On the contrary to the Finno-Ugric
populations, where allele DYS19*14 is the most common
among haplogroup N3 males, allele DYS19*15 is the
most frequent in the Baltic haplogroup N3 (Kasperaviciute
et al. 2004), and was also the most frequent allele in the
haplogroup N3 in northern Belarus (60.0%), providing
additional evidence for the presence of the Baltic substrate
in the genetic pool of Belarusians from the northernmost
part of the country.
Localisation of the Slavic homeland prior to their great
expansion in the fifth to sixth centuries is one of the key

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J Hum Genet (2007) 52:406414

problems of European history in the first millennium AD.

Although it is assumed that prehistorically the original
habitat of Slavs was Asia, from which they migrated in the
third or second millennium BC to populate parts of Eastern
Europe (Encyclopdia Britannica 2006), a debate concerning the European homeland of Slavs seems to remain
unsolved. Because Slavs unequivocally enter the records of
history as late as the sixth century AD, when their expansion in Eastern Europe was already advanced, different
theories concerning the Slavs geographic origin based on
archaeological, anthropological and/or linguistic data have
been formulated. Two such theories have gained the largest
support among the scientists (Schenker 1995), one placing
the cradle of Slavs in the watershed of the Vistula and Oder
rivers (present-day Poland), and the other locating it in the
watershed of the middle Dnieper (present-day Ukraine).
Our results indicate that using the population-of-origin
approach based on the AMOVA, as many as nine
(P > 0.05) or ten (P > 0.01) populations can be traced back
to the lands of present-day Ukraine, including EasternSlavic Russians and Belarusians, Western-Slavic Poles and
Slovaks, and Southern-Slavic Slovenes and Croats. On the
other hand, the Polish population gave insignificant FST
values in pairwise comparisons with only one (i.e. Ukrainians) or three (i.e. Ukrainians, Slovaks, and Lusatians)
populations (P > 0.05 or 0.01, respectively). Moreover, the
Y-STR genetic distance between Poles and Belarusians,
who are geographic neighbours (Table 1), excludes significant gene flow between the two populations and localisation of Belarusians ancestors in present-day Poland.
In conclusion, we have demonstrated that Y-STR haplotype distribution divides Slavs into two genetically distant groups: one encompassing all Western Slavs, Eastern
Slavs, Slovenes and Western Croats, and the other
involving all remaining Southern Slavs. Many northern
Slavic populations are genetically indistinguishable in regard to the nine-locus Y-STR haplotype variation, and this
homogeneity extends from the Alps to the upper Volga,
and even as far as the Pacific Ocean (eastern Russia),
regardless of linguistic, cultural and historical affiliations
of the Slavic ethnicities. The example of Slovaks and Belarusians shows that this homogeneity is likely to be extended to other Y-chromosomal microsatellites as well.
Results of the interpopulation Y-STR haplotype analysis
exclude a significant contribution of ancient tribes inhabiting present-day Poland to the gene pool of Eastern and
Southern Slavs, and suggest that the Slavic expansion
started from present-day Ukraine, thus supporting the
hypothesis that places the earliest known homeland of
Slavs in the basin of the middle Dnieper. To our knowledge, this is the first report on the use of genetic markers in
solving the question of the localisation of the Slavic
homeland.

J Hum Genet (2007) 52:406414

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