UDK 902(497)"633/634":577-2
Documenta PraehistoricaXXXIII (2006)
Most recent investigation of peopling
of Bosnia and Herzegovina: DNA approach
Damir Marjanovic12, Naris Pojskic1, Belma Kalamujic1, Narcisa Bakal1, Sanin Haveric1,
Anja Haveric1, Adaleta Durmic1, Lejla Kovačevic1, Katja Drobnič3,
Rifat Hadžiselimovic1, Dragan Primorac4,5
1	Institute for Genetic Engineering and Biotechnology, University of Sarajevo, Bosnia and Herzegovina
damir.marjanovic@ingeb.ba
2	Center for Integrative Genomics, Molecular Diagnostics, Cell and Gene Therapy, "Rudjer Bošković"
Institute, Zagreb, Croatia
3 Forensic Laboratory and Research Center, Ministry of the Interior, Ljubljana, Slovenia
4 Medical School at Split University, Split, Croatia
5 Medical School at Osijek University, Osijek, Croatia
ABSTRACT - Many historical episodes marked Bosnia and Herzegovina as a significant ethnic cros-
sroads, which makes it a very interesting site for various population studies. The first stages of these
complex investigations were based on observations of numerous phenotype markers. The following
phase, which was relatively brief, was dominated by the use of different cytogenetic markers. Finally,
at the beginning of this century, the molecular-genetic diversity of the BiH population became the
focus of modern research. Autosomal and Y-STR markers, together with mitochondrial haplogroup
(Hg) diversity were initially used in the examination of isolated groups, as well as the whole popu-
lation of modern Bosnia and Herzegovina. The most recent study describes the distribution of Y-chro-
mosome haplogroups in the three main ethnic groups in Bosnia and Herzegovina, and suggests a
preliminary hypothesis for the process of peopling this area.
IZVLEČEK - Veliko zgodovinskih dogodkov je zaznamovalo Bosno in Hercegovino kot pomembno
etnično stičišče, ki je zelo zanimivo za različne populacijske študije. Začetki kompleksnih raziskav
so povezani z opazovanjem številnih fenotipskih označevalcev. V naslednji fazi je prevladala upora-
ba različnih citogenetskih označevalcev. Na začetku tega stoletja so raziskave usmerjene v moleku-
larno-genetsko raznolikost BiH populacije. Pri raziskavah izoliranih populacij, kot tudi celotne člo-
veške populacije moderne Bosne in Hercegovine, so na začetku uporabili avtosomalne in Y-STR oz-
načevalce skupaj z raznolikostjo mitohondrijske haploskupine (Hg). Najnovejša študija opisujepo-
razdelitev haploskupin Y-kromosoma pri treh glavnih etničnih skupinah v Bosni in Hercegovini in
izdela preliminarni scenarij procesa poselitve tega področja.
KEY WORDS - molecular markers; Y-chromosome; Last Glacial Maximum; Paleolithic; Neolithic
Bosnia and Herzegovina: from the most ancient
times to the present
Because of its great number of small, more or less
isolated, indigenous populations, Bosnia-Herzegovina
represents a very interesting area for population-ge-
netic surveys of different levels and approaches. The
fascinating diversity of local human populations (ob-
vious even in phenotype characteristics of individu-
als), as well as various levels of reproductive isola-
tion can be attributed to the simultaneous and inter-
active effects of a vast number of factors (geographi-
cal, ethnic, religious etc.). Within those areas, each
local population exhibits a different degree of open-
ness and isolation and has unique relationships with
ancestral populations. The results of previous studies
show that classic "islands in the land" exist in some
regions (Marjanovic et al. 2004).
Migration played a very important role in shaping
recent conditions, such as, in particular, those rela-
ting to the peopling of central Balkan, as well as
small regional movements, which were rather fre-
quent because of the geographical position of this
area (Malcolm 1994).
Archaeological finds indicate that the territory of mo-
dern Bosnia-Herzegovina was continuously settled
since the Palaeolithic (Imamovic et al. 1998). Stone
tools found in north Bosnia date to more than 50 000
years ago (Imamovic et al. 1998). However, the most
interesting prehistoric traces of civilization date from
the Neolithic.
At the beginning of the second millennium BC, the
discovery of considerable sources of various kinds
of metals attracted new settlers to the Balkan region,
including Bosnia-Herzegovina as its central part. In
this period the area was inhabited by Illyrian tribes.
At the same time, Gals' invasions caused the Illyrians
to migrate and settle in the eastern parts of modern
Bosnia (Wilkes 1992).
By the end of the first century BC, after a hundred-
year war, the Romans had defeated all the Illyrian
tribes. The Illyrian province was under Roman rule
for over 500 years. This was of great importance for
this region that, after the construction of military
roads and opening of numerous mines (silver, gold,
copper, iron etc.), became once again the destination
for many adventurers from all parts of the Roman
Empire (Klaic 1990). The final division of the Ro-
man Empire made the Balkans an unstable border-
land and the crossroads of civilizations, peoples and
religions.
Two crucial, relatively recent events, along with a
number of more minor historic episodes, laid the
foundations for the current population structure of
region.
The first event was the great migration in the sixth
and seventh centuries AD that caused the final fall
of, first, Western then the Eastern Roman Empire.
Documentary evidences suggest that the Goths were
the first immigrants to this region (Malcolm 1994);
shortly thereafter the Avars and Slavs arrived. In
spite of the numerous invaders, there are reasons to
believe that the first invasions were blunted by the
resistance of well-armed resident Illyrians and the
specific landscape, and that the local inhabitants
probably completely assimilated most of the first im-
migrants (Imamovic et al. 1998). According to some
authors, this situation persisted for the next couple
of decades, when the total 'slavization' of this area
began with the arrival of two new Slavic tribes and
the foundation of their powerful neighboring coun-
tries (Imamovic et al. 1998). The present territory
of Bosnia-Herzegovina is located between the areas
inhabited by Croats (to the west) and Serbs (to the
east). The first document mentioning this region
was written by Constantine Porphyrogenitus in 958
(1967). It mentions that Bosnia, being a political and
geographical subject, is located in a narrow area
along the river. This part of history, in the absence
of detailed written documents, is something of a
'twilight zone' for the interpretation of the political
situation of that time. Principally, different under-
standings and presentations of historical facts have
been dictated by ideology and politics. However,
great mysteries such as the Bosnian heretical church,
the term 'Bogomils' (Rački 1931) and the origin of
isolated Bosnian populations remain issues for the
future. A key fact is that the territory of Bosnia-Her-
zegovina at that time was the border, or the cross-
roads of two religious philosophies, Orthodox and
Catholic.
The second important historical moment for this
region was the expansion of the Ottoman Empire
in the fifteen century. The conversion to Islam of a
large part of the population was a fundamental pro-
cess in the creation of modern Bosnian history. It
was a long-term process, which took 150 years be-
fore Muslims were the majority in the former Bo-
snian sandzak (Malcolm 1994). This process is some-
what controversial, and the only thing that can be
refuted with certainty is the hypothesis of forced
mass conversion of the indigenous population. Un-
fortunately, historical dilemmas about some stages
of establishing modern Bosnia-Herzegovina society
have provided opportunities for various ideological,
political, national, even nationalistic manipulations
and distortions of the facts.
Modern Bosnia-Herzegovina is a multi-ethnic and
multi-religious country, with a very stormy recent hi-
story. In various public and scientific media, many
recent conflicts were sought through different inter-
pretations of some parts of its history. Therefore,
every population-genetic study based on the analy-
sis of molecular markers suitable for detecting mi-
gration and micro-evolutionary phenomena, with
utterly reliable and exclusively scientific result pre-
sentation, can create new perspectives, but also raise
questions about the structuring demographic charac-
teristics of this area.
The development of population genetic studies
Fundamental genetic studies of different human, ani-
mal or plant populations are based on a preliminary
understanding of the nature of molecular marker va-
riations in observed populations. This universal rule,
applicable to a wide range of organisms as potential
research subjects, is especially important when Ho-
mo sapiens is the focus of study.
Modern population-genetic studies have changed sig-
nificantly, especially in the last two decades. Previous
studies were based on the observation of classical
phenotypic markers. Population-genetic studies that
simultaneously analyze the diversity of numerous,
carefully selected, phenotypic characteristics can re-
sult in interesting information about micro-changes
in the genetic structure of observed populations over
a certain period. However, many of the informative
phenotypic markers are limited by a lack of clarity
in defining the heritable and non-heritable compo-
nents that form most of the phenotypic attributes of
an organism, especially those which are easiest to
detect.
Human population genetics often uses intra- and in-
ter-population distributions of genetic molecular
markers in and between analyzed parts of popula-
tion in order to elucidate their demographic and mi-
gration history (Wells et al. 2001). The development
of new molecular-biological techniques has improved
the molecular markers system. Among these mar-
kers, of primary importance are the STR (short tan-
dem repeats) sequences, located on autosomal chro-
mosomes (Bowcock et al. 1994), and SNP (single nu-
cleotide polymorphisms) haplotypes (Jin et al. 1999).
Even nowadays, these two kinds of markers can sig-
nificantly contribute to a better understanding of the
demographic and migration processes that affected
the gene pool of now living populations.
Polymorphic markers, such as biallelic and STR, lo-
cated in the non-recombining part of Y-chromosomes
(NRY), are also known as holiandric markers, be-
cause of their transmission from father to son. The
absence of recombination, as well as patrilocal mar-
riage in approximately 70% of modern societies,
makes this unique item of human chromosomal sets
exceedingly useful in the study of large-scale evo-
lutionary events in human history and population
genetics phenomena in large and small, relatively
isolated communities.
In the past few years a substantial amount of pop-
ulation data regarding the non-recombining part of
Y-chromosomes has appeared (YCC 2002 and ref-
erences therein). The spate of surveys in this field
began with papers in the mid-1980's (Casanova et
al. 1985) Markers located in this part of the human
genome have an important role in evolutionary stu-
dies (Hammer et al. 1996; Poloni et al. 1997; Ros-
ser et al. 2000), as well as in forensic (Jobling et al.
1997) and medical genetics (Jobling and Tyler-Smith
2000). The most frequently used markers at that
time were RFLP (restriction fragment length poly-
morphism) or STR loci (FCC 2002). A historic break-
through was made by the utilization of the DHPLC
(denaturing high-performance liquid chromatogra-
phy) method in the investigation of binary markers
from the non-recombining portion of Y-chromo-
somes (NRY) (Underbill et al. 1997). As reported in
some papers (Underbill et al. 2000; Sben et al. 2000;
Hammer et al. 2001), over a short period of time,
this method allowed the identification of over 200
SNPs and short insertion/deletion (indel) positions
(FCC 2002). A very low mutation rate characterizes
these polymorphisms' (named for this reason Unique
Event Polymorphisms - UEP), making them suitable
for the identification and detection of parental line-
ages stretching back thousands of years (FCC 2002).
In order to set up a more flexible nomenclature sys-
tem and to provide a complex NRY binary haplo-
group tree, Y Chromosome Consortium (YCC), estab-
lished by leading experts in Y-chromosome analysis,
suggested a universal system in 2002 that is widely
accepted today (FCC 2002). According to this system,
Y-specific binary haplogroups (often denoted simply
as haplogroups) refer to chromosome clusters charac-
terized by the deepest UEPs. These clusters are more
stable, but at the micro-evolutionary level, less infor-
mative than haplotypes defined by the more vari-
able STR markers (Jobling and Tyler-Smitb 2003).
On the whole, a total of 16 major haplogroups (de-
noted by letters: A-R) were identified and the study
of their distribution (phylogeography) provided im-
portant information about the peopling of different
geographical areas, post-glacial and Neolithic expan-
sions (Underbill et al. 2001). As to Europe, more
than 90% of the Y-chromosomes belong to a few
(E-SRY4064, I-M170, G-M201, J-12f2, R-M17and
R-269) principal haplogroups (Semino et al. 2000;
2004; Cruciani et al. 2004; Rootsi et al. 2004).
Overview of population-genetic studies in Bos-
nia and Herzegovina during the last three cen-
turies
Anthropo-genetic studies in Bosnia-Herzegovina were
begun by the Austro-Hungarian government (Hadži-
selimović 1984). The first known bio-anthropological
analyses of the population conducted by Austro-Hun-
garian army doctors can be considered an initial phase
in determining its genetic structure (Himmel 1887).
It is important to stress that all of the studies were
based primarily on the observation of phenotypic
traits such as blood groups (Kalić, Kostić 1934; Ber-
berović et al. 1973), color blindness (Berberović
1969), the shape of lobules (Hadžiselimović 1970),
PTC sensibility (Berberović et al. 1973), thumb flexi-
bility (Hadžiselimović & Brdar 1979), or analysis of
the effect of some factors that disturb genetic equili-
brium (Hadžiselimović 1983).
The first steps were taken in investigating the variety
of the fundamental, molecular-nuclear and mitochon-
drial DNA markers for diversity analyses of isolated
Bosnia-Herzegovina populations at the beginning of
this century (Marjanović et al. 2004; Kapur et al.
2004; Marjanović et al. 2005). Following analysis of
STR marker variation, autosomal (Marjanović et al.
2006), and Y-chromosome markers (Marjanović et
al. 2005a) were studied in order to establish a refe-
rence sample and to incorporate data about the mo-
lecular biodiversity found in BiH in a regional data-
base (REBID-a).
Finally, the most interesting and controversial re-
sults were obtained trough the observation of 28 Y-
chromosome biallelic markers within 256 males (90
Croats, 81 Serbs and 85 Bosniacs) from Bosnia-Her-
zegovina (Marjanović et al. 2005b; 2005c).
The Y-chromosome approach (summary of pub-
lished articles)
The people of Bosnia-Herzegovina display European
specific haplogroups that probably appeared in dif-
ferent glacial refuge areas of Europe (I-M170, R-
M17 and RM269 from Balkan, Ukrainian and Fran-
co-Cantabrian refuges, respectively), and haplogroups
considered to have originated in Africa (E-SRY4064)
and the Middle East (J-12f2) and to have arrived in
Europe through a prolonged gene flow.
However, the presence of the HgI sub-haplogroup
I-P37 in more than 50% of the examined Y chromo-
somes sets the population of Bosnia-Herzegovina
apart from the majority of other European regions.
Taking into account that a Palaeolithic origin for the
P37 mutation in this Balkan district has been sugges-
ted (Rootsi et al. 2004), it is possible that the post-
glacial expansion of a population with a high fre-
quency of I-P37 from one of the refuges in the Bal-
kans played a major role in the creation of the gene
pool of modern Bosnia-Herzegovina.
HgE is the second most frequent haplogroup in Bo-
snia-Herzegovina; its presence in Europe has been
attributed to multiple migrations from the Middle
East and North Africa during and after the Neolithic
(Cruciani et al. 2004; Semino et al. 2004). HgE is
almost exclusively represented by the sub-clade E-
M78. It is worth mentioning that the clinal distrib-
ution in Europe of E-M78 and its internal micro-sa-
tellite variance have been attributed to dispersals in
Neolithic and post-Neolithic times from the Balkans
to all directions, as far as Iberia to the west and, pro-
bably also to Turkey in the southeast (Cinnioglu et
al. 2004; Cruciani et al. 2004; Semino et al. 2004).
In this framework, our data suggest that this expan-
sion would have significantly affected the gene pool
of Bosnia-Herzegovina.
Haplogroup J is another haplogroup that arrived in
Europe from the Middle East, and its sub-clades pro-
bably marked complex migration processes during
and after the Neolithic period (Cinnioglu et al. 2004;
Di Giacomo et al. 2004; Semino et al. 2004). In the
Bosnia-Herzegovina population this haplogroup, with
almost all of its known sub-clades, is found mainly in
the Bosniacs. The detected J sub-clades are J-M267,
which has been associated with Arab expansion; J-
M92, which suggests genetic links between Anatolia
and southern Italy; J-M67, which is frequent in the
Caucasus; and finally J-M102, which shows frequen-
cy peaks in the southern Balkans and central-south-
ern Italy.
R-M17 is the prevalent sub-haplogroup of Hg R, as
previously observed in other eastern European po-
pulations (Semino et al. 2000; Passarino et al.
2001; Wells et al. 2001). Its frequency perfectly fits
the expected distribution of R-M17, which is found
almost exclusively in Eastern Europe, with a decrea-
sing gradient from northeast to south-west. This gra-
dient, initially attributed to expansion(s) from a Uk-
rainian glacial refuge (Semino et al. 2000), could
also be due to infiltrations of Indo-European speak-
ing peoples from southern Russia about 2000 years
ago (Jovanović 1979; Barać et al. 2003), as well
as to the arrival of Slav clans during the 6th and 7th
centuries. However, to evaluate this latter hypothe-
sis, detailed analyses of R-M17 and internal STR di-
versity in the Bosnia-Herzegovina population are re-
quired.
The presence of the R-M269 sub-haplogroup in the
sample is of interest. Despite the relatively low fre-
quency of this marker, it indicates that the gene pool
of the ancestral population(s) of the Franco-Canta-
brian refuge area also contributed to some extent to
this region of the Balkans, as additionally attested
by analyses of the mtDNA (Achilli et al. 2004).
The first PC analysis (Marjanović et al. 2005c) sug-
gests that the three ethnic groups are genetically ex-
tremely close to each other, and closely related to
other populations of the Balkans. However, the se-
cond PC tends to separate the Croat group from
other populations, probably as a result of the afore-
mentioned effects of genetic drift and founding
events on this ethnic group.
On the whole, our data suggest three main factors
in the gene pool foundation of modern Bosnia-Her-
zegovina: (a) post-LGM expansion - possibly from a
LGM refuge area in the Balkans, (b) complex migra-
tory processes from Central Asia and Eastern Europe
and (c) the numerous Neolithic and Post-Neolithic
migration events that contributed haplogroups con-
sidered to be of African and Middle Eastern origin.
The main objectives of possible future studies
of the peopling of Bosnia and Herzegovina
The outcome of this study is a possible account of
the pattern of settlement in the area of modern Bos-
nia-Herzegovina. For the purpose of further elabora-
tion it is necessary to insist on more of the basic pa-
rameters in future studies.
A multi-disciplinary approach is required in order to
investigate fully all the pre-historic and historical
events relative to the peopling of Bosnia-Herzegovi-
na and the Balkans. All historical, archaeological and
other findings relevant to determining migratory fac-
tors and modern micro-migratory processes should
be analyzed. The results of this study should be com-
pared to those of previous population-genetic studies
based on observations of a wide range of both clas-
sic and molecular markers. Presented scenario, ba-
sed on the results of holandric marker variation ana-
lysis, could benefit from such an approach.
The second parameter, which points to the dating of
primary mutations in samples of the Bosnia-Herze-
govina population, is highly notable. With the inten-
tion of estimating the possible time of mutation oc-
currence, analyses of Y-chromosome STR markers,
and their frequencies and correlations within the
detected haplogroups can be of great value. The re-
sults of these analyses would facilitate the precise
reconstruction of regional migration patterns. Preli-
minary results based on the observation of 100
males from Bosnia and Herzegovina, have already
yielded interesting results (Marjanović et al. 2006).
Eighty-one different Y-STR haplotypes (from a total
number of 100 samples obtained) were detected: 69
of them were unique, 7 appeared twice, 4 appeared
three times, and 1 five times. Six of twelve not single-
ton haplotypes were shared by different population
groups: two of them by Croats and Bosniacs, two by
Bosniacs and Serbs, and the last two by Serbs and
Croats, thus testifying almost certainly to recent gene
flows between groups.
Most importantly, the results must be evaluated with
an expanded sample. The sample of 256 individuals
is larger (in European proportions and for such a re-
latively small population as Bosnia-Herzegovina) than
many other samples investigated so far. Also, sam-
ple size for each of the three major ethnic groups
completely meets the standards for this type of in-
vestigation, and is larger than those used in previ-
ous European studies of much larger populations
than this one. The fact that the subjects are from
more than 50 different locations in Bosnia-Herzego-
vina speaks in favor of a representative sample. How-
ever, regarding the high specificity and close rela-
tions of the ethnic groups, their common historical
development, specific micro-migrational paths and
the specific historical circumstances under which
these groups developed, it is important to examine
the intra-population diversity of regional subpopula-
tions. For instance, the Croat population could be
analyzed including sub-samples from Herzegovina,
Middle Bosnia and Posavina; the Serbs through Her-
zegovina, Eastern Bosnia and Krajina groups; while
the Bosniacs could be typed by comparing Herzego-
vina, East-Middle Bosnia and Western Bosnia groups.
However, all the other demographic-genetic charac-
teristics of the local groups of these populations can
be taken into consideration. This approach can re-
solve the question of whether differences observed
through this study are due to inter-populational (eth-
nical) or, more probably, intra-populational (regio-
nal) characteristics.
Until these complex studies are conducted, it is im-
portant to state that all findings, implications, hypo-
theses, reconstructions and models of this study are
based on the results of the analyses of a screening-
sample of the Bosnia-Herzegovina population and its
relation to previously studied Eurasian populations.
Recent scientific studies from this field (Semino et
al. 2000; Barać et al. 2003, and others) offer one
truly interesting and completely new scenario. How-
ever, extreme caution is required in presenting these
scientific results in the public media in order to avoid
misinterpretation of the facts and unscientific specu-
lation that could have unforeseeable consequences.
-ACKNOWLEDGEMENTS-
All F-cbromosome SNP analyses and results were ob-
tained at tbe Dipartimento di Genetica e Microbiolo-
gia 'A. Buzzati-Traverso', Universit'a diPavia (Pavia,
Italy). Tberefore, we are grateful to Professor Silvana
Santacbiara-Benerecetti, Antonio Torroni and especi-
ally to Ornella Semino and ber associates (Simona,
Silvia, Alessandro, Enza etc.) for tbeir kindness and
support.
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