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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 134:354–368 (2007

)

Comparison of Maternal Lineage and Biogeographic
Analyses of Ancient and Modern Hungarian Populations
Gyöngyvér Tömöry,1,2* Bernadett Csányi,1 Erika Bogácsi-Szabó,1 Tibor Kalmár,1 Ágnes Czibula,1
Aranka Cso†sz,2 Katalin Priskin,1 Balázs Mende,2 Péter Langó,2 C. Stephen Downes,3
and István Raskó1
1
Institute of Genetics, Biological Research Center of the Hungarian Academy of Sciences, 6726 Szeged, Hungary
2
Archaeological Institute of the Hungarian Academy of Sciences, 1014 Budapest, Hungary
3
School of Biomedical Sciences, University of Ulster, Coleraine BT521SA, Northern Ireland

KEY WORDS ancient DNA; mtDNA; Hungarians; Seklers; 10th–11th century bones; Hungarian
conqueror

ABSTRACT The Hungarian language belongs to the genetic effect of populations who came into contact with
Finno-Ugric branch of the Uralic family, but Hungarian ancient Hungarians during their migrations are seen.
speakers have been living in Central Europe for more Strong differences appear when the ancient Hungarian
than 1000 years, surrounded by speakers of unrelated samples are analyzed according to apparent social sta-
Indo-European languages. In order to study the conti- tus, as judged by grave goods. Commoners show a pre-
nuity in maternal lineage between ancient and modern dominance of mtDNA haplotypes and haplogroups (H,
Hungarian populations, polymorphisms in the HVSI R, T), common in west Eurasia, while high-status indi-
and protein coding regions of mitochondrial DNA viduals, presumably conquering Hungarians, show a
sequences of 27 ancient samples (10th–11th centuries), more heterogeneous haplogroup distribution, with hap-
101 modern Hungarian, and 76 modern Hungarian- logroups (N1a, X) which are present at very low fre-
speaking Sekler samples from Transylvania were ana- quencies in modern worldwide populations and are
lyzed. The data were compared with sequences derived absent in recent Hungarian and Sekler populations.
from 57 European and Asian populations, including Modern Hungarian-speaking populations seem to be
Finno-Ugric populations, and statistical analyses were specifically European. Our findings demonstrate that
performed to investigate their genetic relationships. significant genetic differences exist between the ancient
Only 2 of 27 ancient Hungarian samples are unambigu- and recent Hungarian-speaking populations, and no
ously Asian: the rest belong to one of the western Eura- genetic continuity is seen. Am J Phys Anthropol
sian haplogroups, but some Asian affinities, and the 134:354–368, 2007. V 2007 Wiley-Liss, Inc.
C

Hungarian is one of the few non-Indo-European lan- forests and split off from them sometime in the Late
guages in Europe, and the only one in Central Europe Bronze Age or Early Iron Age (around 500 BC); this sep-
(Rédei, 1998). It belongs to the Finno-Ugric branch of aration marks the time from when we can speak of inde-
the Uralic linguistic family, a diverse group of people pendent proto-Hungarian people, the only Finno-Ugric
related by an ancient common linguistic heritage (Hajdú, people to have become horse-riding steppe pastoralists
1976; Napolskikh, 1995). Of the approximately 25 mil- and nomads (Róna-Tas, 1999). Migrating westwards,
lion Finno-Ugrian speakers, three form separate nations: they settled between the Urals and the Middle Volga
the Estonians and Finns on the Eastern Baltic Littoral, region (Fodor, 1976) in a region lying at the Kama-Volga
and the Hungarians (Magyars) in the Danubian plain. confluence, still known to Europeans in the 13th century
Hungarian is also widely spoken by Magyars in western as Magna Hungaria, ‘‘Old Hungary’’ (Fodor, 1982).
Romania, by Seklers (Székelys) in Transylvania, and by Around 700–750 AD the ancient Hungarians left Magna
Csángós east of the Carpathians. Other Finno-Ugric Hungaria and drifted southwards, settling in the steppe
speakers include the Saamis (Lapps) in northern Finno- and forested steppe zone around the river Don, in a
Scandian and Kola Peninsulas, the Erzas, Moksas, region they called Levedia. Here they were neighbors
Maris, Udmurts, and Komis in the northern woodland and allies of the Khazars, a Turkic-speaking people of
zone of European Russia and the Voguls and Ostyaks
around the river Ob in western Siberia (Vékony, 2002).
Grant sponsor: Hungarian National Research and Development
Distantly related to the Finno-Ugrians are the various Programs; Grant numbers: 5/088/2001, 5/038/2004.
Samoyed peoples of Siberia, the Nenets, Enets, Nganas-
sans, and Selkups (Chen et al., 1995). *Correspondence to: Gyöngyvér Tömöry, Archaeological Institute
Hungarians entered central European history as seven of the Hungarian Academy of Sciences, 1014 Budapest, Úri u. 49.,
major Magyar tribes that invaded the Danubian Basin Hungary. E-mail: gtomory@archeo.mta.hu
from across the Carpathians in 895 AD (Regino, 1890).
This was the last in a series of migrations (see Figure 1). Received 19 May 2006; accepted 23 May 2007
At an earlier stage of their history they must have been
far further east, for their closest linguistic affinities are DOI 10.1002/ajpa.20677
with the Voguls and Ostyaks of the forest steppes of Published online 13 July 2007 in Wiley InterScience
western Siberia (Fodor, 1982). The Hungarians left the (www.interscience.wiley.com).

C 2007
V WILEY-LISS, INC.

This migration took the Magyars into the cribed by Kalmar et al. Savirs and Kabars (Kristó. Goths. by treatment with 350 ll before the Magyars’ arrival. 1989). DNA derived from modern samples was ulations are complex. 2004). bones. MATERIALS AND METHODS Sampling Seventy-nine bone samples from ancient remains from the age of the Hungarian conquest were included in the Fig. The whelming majority of the indigenous population was column was washed twice and DNA was eluted in a Slavic. accompanied by the Kabars who had rebelled Sekler individuals living in Romanian Transylvania. from the 10th–11th century.. we have tested the hypothesis that no genetic continuity exists between ancient and modern Hungarian-speaking popu- lations. Kittles et al. 1994). 1. In the early 850s the Hungarians. that they migrated from Hungary later. Samples were excavated in cemeteries from the 10th–11th centuries from different regions of Hungary (Table 1. is still debated. the Hungarian kingdom. Fifteen of the 42 samples did not correspond to (Berta Róna-Tas. 16182–16420). according to the published protocol Hungarian conquest (Bóna. The aim of the present study was to analyze the mitochondrial lineages of human bone samples originating from archaeologically Hungarian graves. The remaining 27 bone samples garian language and culture. 2000). Romans. and Dukla passes across the mountains (Regino. DNA was precipitated from 350-ll extract from bone vania among the mountains. and others: it is 10 min. cur. American Journal of Physical Anthropology—DOI 10. 895 is DNA isolation the traditional date when seven major Magyar and Kabar hordes under King Arpad forced the Verecke. 1990. 1996). spin column and centrifuged at 6000g for 1 min. 1996. (2000) and alternatively when great plain drained by the Danube and Tisza rivers. by Dacians. respectively. ready begun under the Avars. in the Carpathian Basin. and into the more fertile parts of Transyl.. TGATTTCACGGAGGATGGTG-30 (239bp. Huns. under pressure Hair samples were collected from 101 maternally from the Petchenegs. The origin of the Hungarian-speaking Seklers. 1890). 1990). and not entirely understood amplified with the primer pair L16040/H16400. 1999. and from 76 maternally unrelated region. 2000). Around 830 they moved into the eastern Carpathian Mountains. In this modified method. L16201 (50 -AACCCCCTCCCCATGCTTA-30 )/H16400 (50 - The genetic relationships of speakers of the Uralic lan. unrelated modern Hungarian individuals from all ward from Levedia to Etelköz. powder and extraction buffer. 2002. whose steppe subjects and sites and bones were archeologically and anthropomor- allies included the Turkic-speaking Onogur-Bulgars. Estimates of (Walsh et al. Previous analysis of modern Hungarian mitochon- drial sequences indicated an essentially European maternal lineage (Lahermo et al. during GCTTTGGAGTT-GCAGTT-3 ) (240bp. a needed. BIOGEOGRAPHIC ANALYSES OF HUNGARIAN POPULATIONS 355 (Lahermo et al. another Turkish tribe. two partially overlapping that the Seklers were one of the tribes of the original subregions were amplified with the primer pairs L16040 Hungarian conquest. between nucleotide positions 16024–16383. 4M NH4-acetate and 700 ll 96% EtOH at 2708C for matians. Lahermo et al. (Fodor.. (50 -TCTGTTCTTTCATGGGGAAG-30 )/H16239 (50 -GTG 0 others. analysis. 1991). earlier migrants from the DNA isolation from the hair samples was performed Eurasian steppes. and Iranian-speaking we have got PCR results with at least one of the primer Alans from whom the Caucasian Ossetes are descended pairs. against the Khazars (Bálint. the criteria for authentication described below. Fig. In particular. Sar. discarded at this stage. some of whom perhaps lived to see the by using Chelex. 2000).1002/ajpa . phologically well defined before the analysis.. Ancient DNA (aDNA) was isolated from femoral Uzsok. The migrations route of the ancient Hungarians. In 42 cases. 2) and were provided by the Archaeological Institute of the Hungarian Academy of Sciences. moved west. incubated overnight at These regions had been settled for thousands of years 378C (Kalmar et al.. Burial nominally Jewish religion. The mixture was transferred into DNeasy Mini probable on the eve of the Hungarian conquest the over. who settled at the eastern border. Mitochondrial DNA (mtDNA) analysis was performed rently an isolated minority in Romanian Transylvania. the fraction of the total population of the Carpathian Ba- sin consisting of newly arrived Hungarians range from mtDNA analysis: HSVI region 10% to 50% (Cavalli-Sforza. The contact between these peo. on the hypervariable region I (HVSI) of the mtDNA con- which was formerly an autonomous principality under trol region... 16020–16259) and the Middle Ages (Bóna. Slavs. Standard isolation methods were used as des- Pertz. 2002). the Dnieper-Dnester-Prut regions of Hungary. final volume of 40 ll. guages to the neighboring Indo-European-speaking pop. Laitinen et al. Some suggest In cases of ancient samples.. and were ple and the ancient Hungarians is reflected in both Hun. and compare them to samples from modern Hungarians and Seklers. The arrival and settlement of the Slavs has al. a modified method incorporating the DNeasy region that corresponds roughly to present-day Hungary Tissue Kit (Qiagen) was used. were submitted to genetic analysis. Kasperaviciute et al. 1980). Avars. 1998.

38 cycles of denaturation at 938C for 30 s. tomb Hungarian Lowland Early 10th Male anc14 Fadd-Jegeshegy 62. Herrnstadt et al.tomb (10) South-western Hungary Late 10th Female anc15 Fadd-Jegeshegy 63. 1999). The standard amplification reaction contained 4 ll of bone extract. Richards et al. cation conditions were: denaturation at 948C for 6 min. diagnostic polymorphic sites in the HVSII region and cations were purified and concentrated with a Montage mtDNA coding region was also performed. Location of cemeteries where bone samples were was used for cloning of ancient DNA fragments. tomb Hungarian Lowland Late 11th Female anc7 Aldebro†-Mocsáros 25. extension at 728C for 5 min.tomb (3) Hungarian Lowland 10th Female anc4 Szabadkı́gyós-Pálliget 7. 2.tomb South-western Hungary Middle 10th Male anc20 Mözs-Szárazdomb 60. 1996.. 13 PCR buffer and 3 mM mtDNA haplogroups were assigned to each sample by MgCl2 in 40 ll total volume of reaction mixture.5 U AmpliTaq Gold DNA polymerase mtDNA classification (Applied Biosystem).1002/ajpa .tomb South-western Hungary Middle 10th Female anc21 O† rménykút 3/c tomb (12) Hungarian Lowland Late 10th Male anc22 O† rménykút 8. 2001.356 G. after ethidium bromide staining with UV transillumina- tor (UVP). 1999. tomb Hungarian Lowland Late 10th Male anc23 O† rménykút 12.. tomb (6) North-eastern Hungary Late 10th Female anc8 Besenyo†telek-Szo†rhát 1.0 Cycle Sequencing Ready Reaction Kit..tomb (11) South-western Hungary Middle 10th Female anc19 Mözs-Szárazdomb 41. and 30 ll of modern DNA respectively.tomb South-western Hungary 10–11th Male anc18 Mözs-Szárazdomb 2. TÖMÖRY ET AL. TABLE 1. The origin. DNA excavated and the homeland of the Seklers (Seklerland). 2000. 200 lM each of dNTP (Fermentas). according to the manufac- turer’s recommendation. 1998. age and sex of investigated bone samples Sample Origin Estimated age (century) Sex anc1 Izsák-Balázspuszta (1) Hungarian Lowland Middle 10th Male anc2 Magyarhomoróg 120. analysis of the Products of successful and contamination-free amplifi. 2.. Kivisild et al. Macaulay et al. 1.tomb (15) North-western Hungary 11th Male ND: not determined. 25 pmol each of primers. tomb Hungarian Lowland Late 10th Female anc24 Zalavár-Kápolna 270. mtDNA analysis: HVSII and coding regions Sequencing In cases when haplogroup categorization was not pos- sible on the basis of HVSI motifs alone. 1998. and 8 ll Terminator Ready Reaction Mix were mixed in a final volume of 20 ll sequencing reaction.tomb South-western Hungary Late 10th–early 11th Female anc16 Fadd-Jegeshegy 74. The sequences were determined on an ABI Prism 310 sequencer (PE Applied Biosystem). 1993. tomb Hungarian Lowland Late10th Male anc12 Sárrétudvar-Hı́zóföld 118. PCR Centrifuge Filter Device (Millipore) in a final volume of 10 ll in cases of ancient DNA.tomb South-western Hungary Late 10th–early 11th Male anc17 Fadd-Jegeshegy 88. Numbers in brackets refer to cemeteries on the map in Fig. the use of published criteria (Torroni et al. extension at 728C for 40 s. from six selected clones was sequenced in each cases with the universal M13 forward primer (50 -CGCCAG GGTTTTCCCAGTCACGAC-30 ). tomb (7) North-eastern Hungary 10th Male anc9 Eger-Szépasszonyvölgy 4. tomb (8) North-eastern Hungary 10th Nd anc10 Sárrétudvar-Hı́zóföld 5.tomb South-western Hungary Late 10th–early 11th Male anc27 Fadd-Jegeshegy 78.. Amplification reaction was 2004). Ten microliters of purified PCR product. 12. Fin- 568C for 1 min.. tomb (5) Hungarian Lowland Early 11th Female anc6 Szegvár-Oromdu †lo† 593. tomb Hungarian Lowland Middle 10th Female anc13 Sárrétudvar-Hı́zóföld 213. Amplifi... tomb (9) Hungarian Lowland Late10th Male anc11 Sárrétudvar-Hı́zóföld 9. relative to the revised Cambridge Reference checked on 8% native polyacrylamide gel and visualized Sequence (rCRS) (Andrews et al.tomb (4) Hungarian Lowland Middle10th Male anc5 Szegvár-Oromdu †lo† 412. annealing at Quintana-Murci et al. Sequencing reactions were performed using the ABI Prism BigDyeTM Terminator v3. 2001.tomb (13) South-western Hungary 11th-12th Male anc25 Harta-Freifelt 10.tomb (14) Hungarian Lowland Early 10th Female anc26 Lébény-Kaszás 80. Polymorphic American Journal of Physical Anthropology—DOI 10.5 pmol of the same primer which had been used for the amplification.tomb (2) Hungarian Lowland Middle 11th Male anc3 Orosháza-Görbics tanya 2. Richards et al. 2002.. and final nila et al. Cloning of ancient samples The InsT/Aclone PCR Product Cloning Kit (Fermentas) Fig. 1999.

1002/ajpa . Gamma distribution (Corning Incorporated) were used for pipetting. All solutions used were filtered with 0. 2000). T?C L10767 50 -AACCTAAACCTACTCCAATGCTAAA-30 210871MnlI H10965 50 -GTGAGGGGTAGGAGTCAGGTAG-30 11719 Coding reg. All workspa- A 276 bp (np 16090–16365) long control-region of ces (laminar flow surfaces. C?T L10292 50 -CCTTTTACCCCTACCATGAGCC-30 Sequencing H10466 50 -TTTATGTAAATGAGGGGCATTTGG-30 10873 Coding reg. T?C L10146 50 -TGACTACCACAACTCAACGGCT-30 110237HphI H10288 50 -AGGTTAGTTGTTTGTAGGGCTC-30 10310 Coding reg. PCR boxes) and appliances HVSI sequences from 204 Hungarian-speaking individ. Comparisons were The sizes of the resulted fragments were checked on 8% performed by the Blastn program (NCBI). The amplification reaction and conditions were 4. G?A L4521 50 -TACCATCTTTGCAGGCACAC-30 24580NheI H4660 50 -AAGGATTATGGATGCGGTTG-30 7028 Coding reg. Analysis of molecular variance by restriction enzyme cleavage or by sequencing. the primers performed by the ARLEQUIN package. Reduced me- used for amplification and the restriction endonucleases dian network from the sequences of Hungarian-speak- used for detection of polymorphisms are reported in ing populations was also constructed with the Network Table 2. Genetic distances were irradiated with 1. The published carried out wearing the appropriate protective clothing sources of these data are presented in Table 3.. The haplogroup-associated polymorphic sites in the HVSII and coding regions. 1993). ase K) irradiated with UV-C light for 30 min.26 (Meyer et al.000 (Millipore) filter and subsequently (except for protein- (Schneider et al. BIOGEOGRAPHIC ANALYSES OF HUNGARIAN POPULATIONS 357 TABLE 2. Sites with lower mutation rates were the same as described previously for the HVSI sequences.0 J/cm2 UV-C light for 60 min before estimated between all populations as linearized Fst use. T?C H14396 50 -CTCCATCGCTAACCCCACTA-30 Sequencing L14527 50 -TTCTGAATTTTGGGGGAGGT-30 14766 Coding reg. A plus (1) sign indicates the presence while minus (2) sign the absence of recognition site. drills. cell culture dishes.. The Fst values were calcu. contamination with modern DNA.0. con- uals together with the sequences of 7752 individuals tainers) were cleaned with bleach and subsequently was grouped into populations. given greater weight. pincers. face mask and laboratory coats). C?T L14638 50 -ACCCCACAAACCCCATTACT-30 114766MseI H14837 50 -AGGAGTGAGCCGAAGTTTCA-30 a Sites are numbered from the first nucleotid of the recognition sequence. G?A L11674 50 -CAGCCATTCTCATCCAAACC-30 Sequencing H11852 50 -GGGGTAAGGCGAGGTTAGC-30 12308 Coding reg. (gloves. appropriate precautions were taken in order to prevent possible The sequences obtained were compared with previ. C?T L12622 50 -CATCCCTGTAGCATTGTTCG-30 Sequencing H12764 50 -AATTCCTACGCCCTCTCAGC-30 14410 Coding reg. 13 buffer and 3 U enzyme in 50 ll their associated control region motifs in the world-wide final volume.22 lm statistics by the use of ARLEQUIN version 2. the primers used for amplification and the type of detection Nucleotid Detection position Location Substitution Primers of mutationa 72 HVSII T?C L28 50 -CAGGTCTATCACCCTATTAACCA-30 Sequencing H132 50 -GGATGAGGCAGGAATCAAAG-30 73 HVSII A?G L28 50 -CAGGTCTATCACCCTATTAACCA-30 173ApaLI H132 50 -GGATGAGGCAGGAATCAAAG-30 4580 Coding reg. The (AMOVA) of the Hungarian-speaking populations was haplogroup-associated polymorphic sites. preparation (excavation and laboratory work) were sampled from 57 Eurasian populations. Reaction was performed for 2 h at 378C. Through- lated by pairwise comparison using the Tamura–Nei out all manipulations Universal Fit Filter Tips model (Tamura and Nei. Contamination precautions and authentication Database comparison and statistical analysis During each step of sample preparation.1 program. American Journal of Physical Anthropology—DOI 10. The restriction enzyme cleavage reaction mix con. mtDNA database (NCBI. All steps of sample ously published sequences of 7752 Eurasian individuals. sites in the HVSII and mtDNA protein coding regions in two-dimensional scaling (MDS) performed by SPSS were amplified with specific primers and analyzed either package version 5. tubes. A?G L12214 50 -CCCCTTATTTACCGAGAAAGC-30 Sequencing H12398 50 -TTGTTAGGGTTAACGAGGGTGG-30 12705 Coding reg. C?T L6962 50 -TTTTCACCGTAGGTGGCCTG-30 17025AluI H7126 50 -TGAAATGGATTTTGGCGTAGG-30 10238 Coding reg. (pipettes. G?A H10220 50 -GCGTCCCTTTCTCCATAAAA-30 Sequencing L10348 50 -GGCCAGACTTAGGGCTAGGA-30 10400 Coding reg. The distribution of mtDNAs present in the ancient tained 25 ll PCR product from ancient DNA or 10 ll Hungarian population was assessed by searching for from modern DNA. PCR was a 5 0. 1999). native polyacrylamide gel and visualized with an UV transilluminator after ethidium bromide staining. Table 3). The resulting and Eppendorf tubes were sterilized before use by matrix of interpopulation Fst values was summarized autoclaving.

(2000) Englisha 242 Helgason et al. (NCBI Database) Bulgariana 141 Calafell et al. (1996). (2004). TÖMÖRY ET AL. Populations.. (2001) Kareliana 83 Sajantila et al. (1996). (2002) Retoroman 15 Pult et al. (NCBI Database) Croatian 60 Harvey et al. (2000). (2001). Baasner et al.1002/ajpa . Richards et al. Richards et al. (1998) Kirghiz-Lowland 49 Comas et al. (1995) Mongolian 101 Kolman et al. Comas et al. (1998) Azerbaijania 48 Richards et al. The surface of a 2 3 5 cm separate rooms. Passarino et al. (2000) Belarus 51 Belyaeva et al. (1994) Ossetian 197 Kaldma et al. (1998). (2000) Danisha 38 Richards et al. (NCBI Database) Iraqia 116 Richards et al. (1996). (2000). (2004) Evenki 37 Kaessmann et al. (NCBI Database) Slavonic-Russian 88 Markina et al. (1996). (2000) Mari 14 Sajantila et al. (NCBI Database) Albaniana 42 Belledi et al. (2000) Russiana 379 Orekhov et al. (2000) Austriana 99 Parson et al. (1998) Komi 15 Voevoda et al. Malyarchuk et al. (NCBI Database) Swedisha 32 Sajantila et al. (2000) Syriana 69 Richards et al. Richards et al. Bone surfaces. (2002) Saami 114 Sajantila et al. Malyarchuk and Derenko (2001). Richards et al. (1995) Kazakh 82 Yao et al. post-PCR analysis) were carried out in bleach and distilled water. Dubut et al. Bermisheva et al. number of samples of which mtDNA HVSI sequence was used for statistical analysis Number of Populations samples References Adygei 84 Lebedeva et al. (2000) Turkish 28 Calafell et al. (2001). TABLE 3. All steps of work (bone powdering. (2004) Italiana 248 Francalacci et al. (1996). Cali et al. (2000). (NCBI Database) BoscoGurin 13 Pult et al. Shimada et al. Piercy et al. Pfeiffer et al. Lutz et al. (1998). (2001). (2001). (1995). Corte-Real et al. (1999). (1993) Estoniana 149 Sajantila et al. (1994) Romaniana 92 Richards et al. (NCBI Database) Slovakian 128 Metspalu et al. (1994). (1995) Serbian 56 Harvey et al. (1996) Uighur 98 Yao et al. Richards et al.ie/molpopgen/data. Sajantila et al. Tagliabracci et al. (1995). (2002) Finnish 230 Kaessmann et al. Mogentale-Profizi et al. Richards et al. (NCBI Database) Cumanian 11 Bogácsi-Szabó et al. Comas et al. (1998) Kirghiz-Highland 43 Comas et al. Richards et al. (2000). (2000) European-Caucasian 236 Coble et al. Helgason et al. DNA extraction. (2000) Buryat 180 Tajima et al. (1996) North-Ossetiana 106 Richards et al. (1994) Bosnian 163 Harvey et al. 2004). (2003) Georgian 168 Reidla et al. Malyarchuk et al. (2001) Greek-Cretean 185 Villems et al. (2000). (1999) Greek 43 Vernesi et al. Hofmann et al. (2000) Armeniana 191 Richards et al. Dimo-Simonin et al. CEPH Database Galiciana 135 Salas et al. before extraction. (1995) Moksha 20 Sajantila et al. Gonzalez et al. (1996). (1997). Richards et al.tcd.htm (McEvoy et al. (2002) Oberwallis 20 Pult et al. (2000) Basquea 156 Bertranpetit et al. (1998) Ukrainan 17 Malyarchuk and Derenko (2001) Total 7752 a HVSI sequence data were retrieved from the web site www. (NCBI Database) Germana 582 Richards et al. were washed with amplification. part of the bone diaphysis was removed with a sand disk American Journal of Physical Anthropology—DOI 10. (NCBI Database) Palestiniana 117 Di Rienzo and Wilson (1991). McEvoy et al. (1998). (2004). (1996). (2000) Norwegiana 629 Opdal et al. (2000) Polisha 473 Richards et al.358 G. Rousselet and Mangin (1998). (2005) Czecha 83 Richards et al. (1996). (2000). (NCBI Database). (1998). Sajantila et al. (2000) Irisha 300 Richards et al. (1996) Swissa 224 Pult et al. (NCBI Database) Kurdistaniana 53 Richards et al. (1995) Frencha 379 Richards et al. (2001).gen.

9% the ancient bone samples. the In order to detect possible contamination by exogenous ancient Hungarian frequency is lower than in modern DNA. which was homologous with the Some of the ancient sequences (anc4. rect and the ancient samples are free of contamination. I. However. Polymorphisms in ancient Hungarian samples Sample HVSI haplotypes (216000)a HVSII and coding region haplotypesa Haplogroup anc1* 140C 182C 183C 189C 217C 274A 9bp del B anc25* 294T 304C 273ApaLI 27025AluI 214766MseI 10310G H anc2 CRS 273ApaLI 27025AluI 214766MseI H anc5 CRS 273ApaLI 27025AluI 114766MseI H anc17 304C 273ApaLI 27025AluI 214766MseI H anc19 093C 366T 273ApaLI 27025AluI 214766MseI H anc21* CRS 273ApaLI 27025AluI 214766MseI H anc26 311C 362C 273ApaLI 27025AluI 214766MseI H anc27 311C 273ApaLI 214766MseI 17025AluI HV anc10 129A 148T 223T 110237HphI 12705T I anc20 223T 311C 10400T M anc3* 147A 172C 183C 189C 223T 320T 355T 110237HphI 10400C N1a anc8* 147A 172C 183C 189C 223T 248T 320T 355T 110237HphI 110871MnlI N1a anc14 CRS 173ApaLI 27025AluI 11719A 12308A 114766MseI R anc13* 311C 173ApaLI 11719A 12308A 12705C 114766MseI R anc12* 126C 182C 183C 189C 294T 296T 298C T anc16 051G 126C 147T 294T 296T 10400C 110871MnlI T anc18 126C 148T 218T 294T 304C T2 anc11 126C 294T 304C T2 anc9* 259A 311C 273ApaLI 17025AluI 12308G 114766MseI U anc24 311C 343G 12308G U3 anc7* 356C 12308G U4 anc4* 223T 356C 10400C 12308G 12705C 173ApaLI U4 anc6 114A 192T 256T 270T 294T 12308G U5a1 anc15 153A 298C 72C 273ApaLI 14580NheI 17025AluI 214766MseI V anc22* 183C 189C 223T 278T 110871MnlI 14470C X anc23* 183C 189C 223T 278T 110871MnlI 14470C X a Sites are numbered according to revised CRS (Andrews et al. Two samples (anc22. two independent ian bones were analyzed for mtDNA polymorphisms.(L15561 50 -CACCATACCCACCTGACATGCA-30 and H15741 50 -GCTGATTTCCCGCGGCTTGGTG-30 . buried with human sample maternally related. those two individuals could be burial site at Harta-Freifelt.4% and all persons involved in processing the samples were 15. was isolated with the same procedure as the the subsequent statistical analysis to avoid the bias of human aDNA. respectively. Haplotypes of logroups U and T were represented with 19. and at least two PCR sequences could be assigned to definitive haplogroups. belonging to hap- horse burial (only the horse skull and the first two legs logroup X. The most frequent (26. assess the reproducibility and authenticity of results. Each Hungarian samples bone sample was powdered by at least two scientists. posses the same haplotype and were derived are buried with the human remains). respectively (Table ered authentic only if partial sequences were assembled 5). Only the horse-specific primers resulted in amplification product.9%) was haplogroup H. anc22. To prove the authenticity of each) and one each to HV. to Twenty-five different haplotypes were distinguished.9%). at Twenty-seven sequences derived from ancient Hungar- least two times each. 225 bp (15539–15763)) and human-specific primers (L16040/ Cloning H16239).. haplogroups B (3.2% frequencies. Only one of them was included in anc25. which could be assigned to 15 different haplogroups. anc25) were cloned. Horse aDNA was amplified with both inbred samples. Bone powder was pro- duced by boring. American Journal of Physical Anthropology—DOI 10. excavated from the from the same burial site. These polymorphisms in ancient sequences are shown in ces were sequenced from both directions.9%) and M (3. The others were assigned to European-specific hap- unambiguously. All DNA extractions were carried out. The whole 360 Table 4. and preV haplogroups (3. previously published horse sequences. * Sequences belonging to the classical conquerors. Hap- powder) were used as negative controls. the commonest European haplogroup. 1998). BIOGEOGRAPHIC ANALYSES OF HUNGARIAN POPULATIONS 359 TABLE 4.1002/ajpa . Two samples were determined and compared with the results obtained from assigned to each of the N1a and R haplogroups (7. In each case. in order it is possible to isolate aDNA with this method. Richards. aDNA from a fractional each). bp sequence was assembled from the four independently Two sequences (anc1. to 3–4-mm depth in order to eliminate possible surface RESULTS contamination. horse. extraction and amplification blanks (with no bone Europe (40–60%. Sequences were consid. The cleaned surface was irradiated with Haplogroup identification of ancient UV light at 1.9% ancient human DNA further.0 J/cm2 for 30 min. in average 46%. anc23. This proves that anc12. anc20) belong to typical Asian produced part of the sequence. free of to prove that the results of the direct sequencing are cor- exogenous contamination. and if both collaborators reproduced at logroups. anc6. amplifications were performed from each extract. anc23). Both subregions of the analyzed mtDNA HVSI sequen. least one partial sequence. anc8. anc15. and collected into a sterile tube. 1999).

7 X 3. In case of fragments anc6.7 3. Szo†ke.1 C 2.360 G.0 2.1 U3 2. The N haplogroup is of Status comparison of the two groups of Near Eastern origin (Richards et al. anc12. respectively (9. How.7 3. the sequence of the 240 bp unambiguously typical of classical Hungarian conquer- and the 239 bp long portion of the HVSI region was ors—horse’s skull. mounted belts. and X.0 1.9 T2 4.6 M 3. As the result of cloning.1 6. one of the six clones shows C in the np16223.0 2.6 18.8 W 7.0 U5a 2.2 HV 3. 1962.7 7. 16296.7 N1a 7. 16189.7 K 7.1% each). horse harness.6 9. These are bur- anc25 (240 bp fragments) all the six cloned sequences ial sites with many tombs. anc12.0 1.4 U 4. TÖMÖRY ET AL.9 6.7 V 4.0 preV 1. and it also bears because of the small sample size must be considered 12705C.6 18. this statement ing sample anc4 displays 16223T.. 16192. and decorations. The frequency of haplogroup H is rather low compared with the average in Europe. simpler. 2000).0 1. were excavated from cemeteries with rich grave goods.4 U5b1 1.3 T3 1. 1991). The ancient samples originated from burial sites of the This haplogroup is present in the Carpathian Basin 10th–11th centuries. and U4 (18.7 3.1002/ajpa . 1991).9 26.0 J1a 1.9 6. anc25) arrow-or spear-heads.0 1. anc15.. logroup B (9.9 5. with poor archaeological found between cloned and direct sequences of ancient remains (no horse harness. T.0 2.7 T1 2. ferences were found (Table 4). ing. paring these two groups on the basis of haplogroup fre- respectively) could be defined unambiguously from the quencies and haplotype identities significant genetic dif- sequences of the clones. Bálint.9 6. earrings (Mesterházy.9 6.7 I 2. 1997. fewer.6 9.4 3.0 2.2% each) and one of which raises the possibility of heteroplasmy at this haplogroups R. 2005).3 3.7 H 39. typically those of plebeians: were identical to that obtained by direct sequencing (not who could be either Hungarian invaders or members of shown). According to the direct sequenc.0 U5a1 2. and four cases (anc4.2 33.7 J 5. at about 0.0 J2 1. TABLE 5. However. anc12.9 9. anc3 and anc8 show could be divided into two major groups. Twelve of them 16189C which is characteristic of the Central Asian American Journal of Physical Anthropology—DOI 10.6 13.4 JT 1. and nps 16183. anc22) respectively. since 12705T also splits off the 16223C cluster (Mac.7 U5a1a 2. However. anc23.9 9. established in five (anc6.1 6. Two sequences each are members of ever. These samples In addition in case of anc4 the sequence containing the came mostly from the cemeteries of nuclear families 12705 position was determined after cloning. The fre- ancient samples quency of N1a mtDNA lineage today is very low any- where in the world.0 2. braid ornaments. The cloned sequences confirmed the sequence belongs to the typical Asian-specific hap- coexistence of these mutations in the same sample.6 36. and poorer samples (Table 6). and grave goods) (Szo†ke.7 U4 4.0 3. U.3 J1 1. (Révész. haplogroups H. N1a. position. In case of fragments anc12 (239 bp) and anc23 other populations who had been in the Carpathian Basin (240 bp) obscurities in the directly sequenced samples before the Hungarian conquest (Mesterházy. These two mutations contradict to each other.2 R 7.1%). anc8.4 7.7 7. The bone findings archaeologically from the Neolithic. Haplogroup frequencies in Hungarian-speaking populations in percentages Recent Ancient Classical Haplogroups Hungarian Sekler Hungarian conquerors Commoners B 3. 1962).9 10.9 2. with utmost cautions.9 9. Among the classical Hungarian conquerors one aulay et al.0 7..2 U5 1. The other 15 bone samples originated Only sporadic and inconsequential differences were from commoners’ cemeteries.9 6.1 Total 100 100 100 100 100 and to eliminate ambiguities obtained by direct sequenc.4 T4 1. 1999). Com- (at nps 16294.0 9.7 T 2.9 6.2% (Haak et al. 1996).9 18.

. anc12-direct . . . . T . . . . . T . . . . . . T . . A . . . . . . T T C . . . . anc15-clone4 . . . Sequences of cloned PCR fragments compared to the Cambridge Reference Sequence and the results of direct sequencinga 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 6 0 0 0 0 0 0 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 3 3 4 4 5 7 5 8 8 9 1 2 4 5 6 6 7 9 9 9 1 2 4 5 5 5 6 7 7 7 5 9 8 9 4 4 3 3 9 2 0 3 8 9 0 7 8 4 6 8 9 0 4 5 6 8 5 1 6 8 CRS G G G G A A G A T C A C C C C C C C C T G C C C T C C A C C anc4-direct . . T . . T T . . . . T . . anc8-direct . A . . . . . . . . . anc22-clone4 . . anc8-clone2 . T . . . . . . . . . . R . . . T . . . . . . . . . . A . . . . . . . . . . . . . G . . . . anc23-clone4 . . . . anc23-clone2 . . . . . . . . . anc15-clone3 . . . T . . T T . . C . . . . . . . . . . T . . . . . . anc12-clone1 . . . . . . T anc22-clone3 . . . . Y Y T . . . C C C . . . . . . . . . . . . . A . T . . . anc4-clone4 . . . anc4-clone6 . anc12-clone4 . . . . . . anc22-clone5 . . . . . . . anc8-clone5 . . C . . . A . . . . . . . . . . C T . T . . . . . . . . . . . . . . . . . . . . . . . anc23-clone5 . . . anc12-clone3 . T T . . . . . . . . . . . . . . . . . . TABLE 6. . . . C C C . . A . T . . T . . . . . . . . . . . T . C . . . . . . . T T C . . . C . . . . . . . . . . . . . . . T T C . . . T . anc22-clone6 . . . . . . . . . . . . . . . . . . . T . . . . C . . T . . . . anc22-direct . . . C C C . . . . A . . . . . . . . . . . . . . . anc23-clone3 . . . . . . . . . . . T T . . . anc12-clone2 . anc15-clone6 . . . . T . . A . . . . . G T . . . . . T anc8-clone4 . . anc12-clone5 . . . C . . . T T . . . . . . M Y Y . . . . . T . . A . A . . . . . . . . . . . . . . . . . . . T . . . . . . T . . . . anc22-clone1 . . . . . . . . . . . anc15-clone1 A . . . . . T . . . . . . . . . . . . . . . anc22-clone2 . . . . . . . . C C C . . . . . . anc8-clone3 . . T . . . . T . anc8-clone6 . . . . . anc15-clone5 . . . . . . . . a Only variable sites are shown. . . . . . anc4-clone2 . C . T . . . . . . A . . . . . . . . . . . . T . . . . . anc23-clone6 . . . A A . anc4-clone1 . . A . anc4-clone5 . . T . . . . . . T . . . . . T . . . . . . C C C . . anc12-clone6 . . . . . anc23-direct . . . . . . T T T . . . anc15-clone2 . . T T C . . . . T T C . . T . . . . . . . T T C . . C C C . . . . . . anc8-clone1 . . . . . . . . . . . G . C . . T . . . . A . . . . . . . . . . . . . . . T . . . . . . . anc23-clone1 . anc15-direct . . . . . . . . . . . . . . . . . T . . . . . . . . . . . T . . . . T . T . . . . anc4-clone3 . . . . . . . . . . . T . Dots mean the identical nucleotid with the CRS. A . . . . . . . . . . . . . T T . .

1999. iance between the commoners and modern Hungarians Norwegian. Russian. 1999) cient samples show the lowest genetic distance from originate from commoners’ cemeteries. 0.65% between the which is characteristic of northeastern Europe (Pericic classical conquerors and modern Hungarians. Anc3 is to one of the older (anc11. eight mod. sidering the modern populations the percentage of var- cific haplogroup M (Macaulay et al. Bosnian. studied are concentrated in a group composed of mainly ards et al. ulations under consideration were summarized by a All ancient samples show significant distances (P < reduced median network. and 1. The frequency of haplogroup H is 33. 4a. Eastern. Swiss). a member of haplogroup I. nian. Ukrainian). The two ancient populations were in- Anc10. English. shares haplotypes dependently compared with the modern Hungarian- found in Korean. This hap. and also the Finno-Ugrian Komi. U3. Among the recent Sekler Balkans. Palestinian).3%). A group of Central Asian populations split off American Journal of Physical Anthropology—DOI 10. and Sekler samples is insignificant. It seems that Seklers. while haplogroups HV. modern individuals analyzed in this study.. types. which appears occur in recent Hungarian and 12% in Sekler popula- with high frequency in populations of the Volga. but with some from Central and Western sequences two are members of the Asian-specific Europe.26%) while et al.3%). one (anc20) was assigned to the Asian-spe. While there was a variance of 12. while 23% of ancient haplotypes ces were assigned to the U4 haplogroup.005) to All of the recent Hungarian sequences are members of 25 populations. they appear in all branches together with distance between the commoners and recent Hungarian the other sequences but mostly show individual haplo.05) from the recent Hungarian-speaking populations. (Bandelt. French. Three of the four sequences belonging to the Eu. European populations. Our data were compared with 7752 sequences from logroups H. Spanish. Seven of the 15 sequences belong to the hap. apart from CRS sequences. variance was observed. Swedish. Azerbaijani. and West- The sequences of ancient samples were compared with ern Europe (Czech. compared to the other populations. 1999.62% variance was among the two groups and 2. group have a relatively low distance (Fst < 0.38% of which approaches that in modern European populations. I. Statistical analysis nian populations. Two sequen.. anc27) haplo- at np16248 one step neighbor of Central Asian sequen.9%) are Central Asian (Kazakh. recent samples from 101 Hungarian and 76 Sekler indi. the CRS which is common in Europe (Rich. 2004). respectively. Slovakian. but while the percentage in modern Hun. Moksha. Serbian. types. Slovakian. Con- cemeteries. mostly from Eastern Europe and the European type haplogroups. speaking populations (taking as one group) at variance The Anc6 sequence belongs to the U5a1 haplogroup.65% and 36. the Middle East. branch. U5a1. When the two modern and the The T2 haplogroup is represented by two samples two ancient populations were grouped and related (13. Czech.3%.95% and 18. Kirghiz.005) to populations from Central. while Seklers have very low distances (Fst < 0. see Fig. which are most common in 57 populations from Europe. Estonian.b). HV. and W haplogroups jani. some populations from the Levant (Syrian. 2005). while one share the same HVSI but ian conquerors the frequency of haplogroup U* was high not RFLP haplotype with anc13 and anc27 sequences.75%). Mongolian). Syr- similar to that in Europe among ancient samples and in ian. When the samples of classical Hungarian con- haplogroups. Fst values were visualized by the use of multidimen- As far as the comparison with ancient samples is sional scaling (MDS) (Fig. French. So these haplotypes might have come through Asia CRS sequences only one has an older equivalent in to the Carpathian Basin. This haplotype is shared between Finnish. Ural tions. nated from a Buryat tribe and is a one step neighbor (at only five of the modern haplotypes can be traced back np16183) of other Central Asian type sequences. Moksha).001) from populations from Asia (Turkish. TÖMÖRY ET AL. Haplogroup H is the most frequent in all querors and commoners were analyzed. and R. 0. the former populations.05) from garian-speaking populations (39. in case of the Hungarians. Croatian. Polish. and Oberwallis populations.. The Anc4 sample belonging to the U4 haplogroup has equivalent sequences in the Siberian Nenets. 2000). Ukrainian.2%. (27. and Ossetian populations. R. Central Asia (Turkish. Austrian. On the basis of Fst values (Table 8) the an- ropean-specific T* haplogroups (Macaulay et al. The and Seklers. Swiss. Bos- Haplotype variations among Hungarian-speaking pop.1002/ajpa . and haplogroup C. The ancient Hungarian samples are not separated from as do classical Hungarian conquerors from the com- the other sequences derived from modern Hungarians moners and recent Hungarian-speaking populations. Among the recent Sekler sequences.31% preV are represented by one sequence only (6.. apart from ces.362 G. tral Asia. viduals. the var- et al. sequences. Ukrainian. Among the classical Hungar. Swedish. the others belong to western Eurasian Mari. Serbian. T. Croatian. On the whole 13% of recent Hungarian logroup occurs mainly in the southern but also in the and 9% of the Sekler haplotypes can be traced back to eastern region of Europe (Richards. region. 3). and Georgian is negligible. The modern Hungar- Hungarian-speaking populations ian population shows very low genetic distances (Fst < 0. level. within the groups. anc10 haplotype. Georgian. including recent Hungarians and ern Hungarian and six Sekler samples. Moksha) and also from Europe while the N1a and X haplogroups are missing among (Italian. and Esto.. K. Estonian. ancient haplotypes. Adygei. and Cen- Europe. AMOVA was performed with different grouping of the Among those samples who originated from commoners’ studied Hungarian-speaking populations (Table 7). Richards iance between populations is very low (0. and Comparison of ancient samples with two current Europe (Oberwallis. Most of the populations concerned. Greek). Turk. 1998). while samples from particular among samples from the classical conquerors commoners’ cemeteries show very low distances (Fst < this value is rather low (26. Austrian. Azerbai- The ancient sequences lack the J. Anc8 is identical with only one sequence origi. comparing the two groups of ancient samples 12. anc17. respectively). Finnish. and West Siberia (Malyarchuk. anc25. Slovakian. 2000) was found in two ancient.

mapped with Adygei. Class: Classical conquerors. Ancient 2.0048) 87. * Indicates the Cambridge Reference Sequence.91 (0.229) 97.06 (0. The amount of circles indicates the number of samples with the given hap- lotypes. and the Finno-Ugric under consideration. Uighur. According to our results. and to the mediaeval Cumanian population that migrated into DISCUSSION Hungary in the 13th century. from Central Asia. including the modern Hungarians and Seklers. querors are well separated from all the populations ian.000) Hun1Sek vs.338) 0. 3. Coding region and HVSII sites which define main branches of haplogroups are bold. Comm 21. Comm: Commoners.007) 96. Ukrainian.234) 99. at the levels of mitochondrial haplogroup and haplotype. the ancient Hungarian popu- Kazakh. Sek: Sekler. and it is close to the Azerbaijani. Turkish.62 (0.21 (0.38 (0.655) 0. and Euro. Analyzing the samples of classical conquerors and pean Caucasians are well separated from all studied commoners separately. Kurdish. Kirghiz) and western Eurasian popula. American Journal of Physical Anthropology—DOI 10.234) Ancient (Class vs.326) 0. Saamis. Sek) 0. BIOGEOGRAPHIC ANALYSES OF HUNGARIAN POPULATIONS 363 Fig. Oberwallis populations.0137) Hun1Sek vs. Evenkis.0048) Hun1Sek vs. AMOVA in samples of Hungarian-speaking populations Among Among populations Within Groupsa groupsb within groups populations Modern (Hun vs.26 (0. This group further was mapped between a group of Central Asian (Mongolian.172) 100. The classical con- mapped within a group that includes Palestinian.26 (0.26 (0. Serbian. Class 12.62 (0.62 (0.165) 87.74 (0. b P values are given in parentheses.000) Hun1Sek vs. Comm) 12. unambiguously from them.34 (0. the commoners’ samples are populations. Reduced median network of the haplotypes identified among ancient Hungarian and modern Hungarian-speaking popu- lations. and Ossetian populations The group of the ancient Hungarian samples is close to Western Eurasian populations. TABLE 7. lation from the 10th–11th centuries was heterogeneous tions.332) 2. Class1Comm 1.1002/ajpa . Syr. but they are closest to populations Komi populations.17 (0.416) a Hun: Modern Hungarian.65 (0.10 (0.31 (0. North-Ossetian.

00757 20.10080 Komi 0. T) common in west Eurasia.06093 20.01162 0.01842 0.00000 0.03138 0.16259 0.00794 Belarus 0.04771 0.01449 0.00422 0.12111 20.00124 Polish 0.14113 0.01587 0.01116 0. R.00444 Armenian 0.00315 Adygei 0.10747 20.00275 0.16809 Serbian 0.00251 Azerbaijani 0.25371 0.14129 0.02540 0.00295 20.06311 0.11208 20.07472 0.00771 Palestinian 0.02460 0.07059 0.00520 0.13064 0.00378 0.01856 0.00462 European-Caucasian 0.07145 0.00339 Glacial 0.03327 Swiss 0.02521 Evenki 0.02076 0.01335 Danish 0.01993 0.00785 0.10703 20. TABLE 8.07693 0.00301 Sweden 20.11704 20.11508 0.02338 Retoroman 0.01858 0.04670 0.02943 0.00346 0.02680 0.00013 0.00724 0.02771 0.04438 0.13606 0.00704 Bulgarian 0.06776 0.00932 Mari 0.12231 20.00833 0.10787 20.18442 0.1002/ajpa .00919 0.07162 0.00254 0.00801 Irish 0.07655 0.13045 0.02186 0.09689 20.00000 0. while classi- European haplogroups.08971 20.00834 0.01184 0.05174 0.07410 20.07064 0.00753 0.01767 0.03060 0.01129 0.00348 0.01290 0.05911 0. Some could be classified to haplogroups lations who came into contact with ancient Hungarians (N1a.01276 20.10033 Fifteen haplogroups and 25 haplotypes were distin.02135 Cumanian 20.07671 20.01115 0.00463 0.00488 Uighur 0.11010 20.00511 20.00974 0.01050 Classicals 0. TÖMÖRY ET AL.01872 0. we de.01211 Slovakian 0.09060 0.00486 0.00942 0.00414 20.06779 0.01951 0.01831 0.00662 Sekler 0.00126 Russian 0.02410 Ossetian 0.00330 0.02851 0.02876 Georgian 0.00668 0.01358 0.00376 0.00641 0.11421 0.00257 0.10602 0.01374 0.00032 0.01476 0.01492 0.00002 0.00462 20.04144 0.00382 0.06651 20.01996 20.01038 0.00112 0.00236 Austrian 0.01675 0.12108 20.01475 North-Ossetian 0.00198 0.11408 20.02132 0.01798 0.02561 English 0.03228 Kirghiz-High 0.04939 Ukrainian 20.02401 0.00850 0. Analyzing the haplotypes.13190 20.00566 0.01533 20.10941 0.00322 Estonian 0.01323 0.24119 0.00611 20.00728 0.04968 Kirghiz-Low 0.01420 0.01070 Turkish 0.09332 0.00651 20.09912 0. all but moners show a clear dominance of haplotypes and hap- two (anc1.11356 20.00262 0.07063 0. ous when analyzed according to the burial status: Com- guished among the 27 samples studied.01631 0.04311 0.01573 0.18491 0.00684 BoscoGurin 0. However.01138 0.03066 0.10251 0.05304 0.00063 Karelian 0.01542 0.01860 0.00644 0.01270 Norwegian 0. anc20) ancient samples belong to potentially logroups (H.00185 0.00940 0. The difference is more conspicu. cal conquerors show a more heterogeneous haplogroup tect some Asian affinities.01081 0.00629 Buryat 0.02858 0.01266 0.06349 0.10213 0.00129 0.01377 Moksha 0.02587 0.06363 0.13755 0.01556 0. and are absent in recent Hungarian American Journal of Physical Anthropology—DOI 10. wide populations.00638 0.00392 0.00192 Greek 0.03695 0.00813 0.01164 0.01698 0.00022 0.01191 0.03129 0.01558 20.01146 0.06241 20.05399 20.08331 20.13345 0.00630 Mongolian 0.00607 20.00576 Romanian 0.03161 0.05535 Italian 0.00344 0. Fst values calculated from mtDNA HVSI sequence data Modern Ancient Classical Populations Hungarian Sekler Hungarian conquerors Commoners Iraqi 0.00390 Greek-Cretan 0.11083 Croatian 20.01210 20.00414 0.03570 0.02273 0.14186 0.02032 0.01513 0.03038 0.08389 0.00542 0.00340 0.09576 0.00177 0.03682 Saami 0.08103 0.01578 Modern Hungarian 0.11565 20.00352 Albanian 0.01136 0.00152 German 0.00515 0.06618 0. X) which frequencies are very low in recent world- during their movement.07801 0.00034 20.00676 Kurdistanian 0.02065 20.11701 0.01096 0.00698 French 20.00321 0.01840 0.00452 0.00332 20.01407 0.04136 0.03660 0.18765 0.17773 0.00060 0.00462 20.00107 0.00967 0.01481 0.00347 Czech 20. representation.10656 0.00274 Slavonic-Russian 0.16310 0.00460 0.07964 0.00419 Kazakh 0.01690 0.00361 20.02898 Oberwallis 0.12206 0.00194 20.00612 0.00339 0.02124 0.06995 Bosnian 0.01102 0.22219 0.11320 0.00146 0.20320 Finn 0.05762 0.364 G. and the genetic effect of popu.01535 0.02469 0.00502 0.00876 0.00202 Syrian 0.02424 0.00070 0.10033 0.06400 0.13972 0.01142 Basque 0.

Not only were they in contact there with tions seem to be specifically European populations. modern Hungary.1002/ajpa . The explanation could be that the in the Hungarian conqueror population unambiguously. Reto 5 Retoroman. Gal 5 Galician. and Iranian-speak- Turkish-speaking populations of the Altai (Derenko ing Alans—seems to have left imprints in the ancient et al. Eng 5 English. Bel 5 Belgian. Mok 5 Moksha. Ger 5 German. Russ 5 Russian. Slo 5 Slovakian. It 5 Italian. Swe 5 Swedish. Nor 5 Nor- wagian. 2002. Ady 5 Adygei. misheva et al. 2004). Ir 5 Irish. Dan 5 Danish. they can be placed between Asian and European popula- mate the proportion of Hungarian invaders whose tions closest to Turks. across the Eurasian steppe. as do even the Petchenegs. Onogur-Bulgars. Cum 5 ancient Cumanian. Azer 5 Azerbaijani. and Sekler populations. Bul 5 Bulgarian. Finn 5 Finnish. Ob 5 Oberwallis. BIOGEOGRAPHIC ANALYSES OF HUNGARIAN POPULATIONS 365 Fig. Sw 5 Swiss. lations from Central Asia. over the last few centuries. commoners’ cemeteries might contain the remains of Genetic distances suggest a clear relationship with popu- pre-Hungarian populations. a: the localization of commoners and classical conquerors b: the localization of ancient Hungarian and modern Hungarian speaking populations. (2000). By contrast.. Turk 5 Turkish. Population labels are as follows: Irq 5 Iraqi. Au 5 Austrian. Ukrainians. N-Oss 5 North-Ossetian. Kaz 5 Kazakh. Cre 5 Cre- tean. which can be traced back for over American Journal of Physical Anthropology—DOI 10. was in Komis. recent Hungarian-speaking popula- the steppes. Rom 5 Romanian. Fr 5 French. Ukr 5 Ukrainan. Oss 5 Ossetian. but recent cordance with the data of Lahermo et al. Pol 5 Polish. Comm 5 Commoners. over many millennia. as well as in Hungarian language one in a most complex series of population movement and culture. Est 5 Estonian. The Magyar invasion of Europe was only Hungarian gene pool. Cze 5 Czech. Classicals 5 classical Conqueror. Malyarchuk. Cro 5 Croatian. Sy 5 Syrian. and the Finno-Ugric maternal ancestry. the linguistic and cultural inheritance of directions. in ac- Indo-European speakers such as the Alans. Hun 5 recent Hungarian. Ancient 5 ancient Hungarians. MDS plot of interpopulation Fst values. Kar 5 Karelian. Serb 5 Serbian. Arm 5 Armenian. in all However. Sek 5 Sekler. Savirs.. Sl-Russ 5 Slavonic- Russian. Geo 5 Georgian. Bel 5 Belarus. On a distance matrix tree This level of analysis may very probably underesti. Statistical analysis clearly Statistical analysis shows the Asian genetic influence reveals this difference. Gr 5 Greek. 4. analysis has shown that the mitochondrial characteristics The genetic effect of populations who lived in close of the Finno-Ugric populations of the Volga and Ural contact with the Hungarians during their migration regions have significant west Eurasian elements (Ber. Kurd 5 Kurdistanian. Bos 5 Bosnian. from the Ural region to the Carpathian Basin—Khazars. Pal 5 Palestinian. 2003).

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Genetic diversity in the Iberian Peninsula determined DQ246412–DQ246437. only a small fraction of the total population of the Car. who mixed with other populations had been Belledi M. Raskó I. Mateu E. Coble MD. Czibula Á. logica 11. Maternal and paternal line- Dacians. AF487612–AF487614. 2003. AF487580. 2004. Villems R. Ruhlen M.nlm. Di Rienzo A. Macaulay VA. Forensic evaluation of mtDNA in a popula- Cambridge reference sequence for human mitochondrial DNA. Excoffier L. Richards MB. 2001. AF487596. 1998. Komi). Acta Orient Acad Scientiarum Hungaricae 55:43–67. This Bóna I. Underhill P. EF646857]). Jahrhundert. mtDNA control region and RFLP data for Sicily and France. Germans. pest: MTA Történettudományi Intézete. This is not surprising. population data and maternal inheritance. Tolun A. commoners’ cemeteries and the recent Hungarian-speak. who show the lowest genetic distance populations. A magyarok és Európa a 9–10. There is little direct genetic relationship between the 98:169–178. AF487590. Int J Legal Med 113:89–97. Calafell F. . De Leo G. Tambets K. AF487561–AF487570. Andrews RM. 1991. In the recent Cavalli-Sforza LL. 1991. 1995. the gene pool. since during the ian. Issad AF487574–AF487576. 1000 years. Ann Hum Genet 60: has not lead to significant genetic isolation.). Worldwide analysis of the Seklers the genetic effect of Eastern and Southern genetic and linguistic relationships of human populations. Leho†cz for skilled technical assistance.gov/Genbank/index. Romans. Genetic analysis of single hair shafts by auto- ACKNOWLEDGMENTS mated sequence analysis of the mitochondrial d-loop region. Comas D. Nat Genet 23:147. [Diversity of mitochondrial DNA haplotypes in ethnic popula- The classical Hungarian conquerors and the modern tions of the Volga-Ural region of Russia]. Hum Biol 67:595–612. Europeans is more visible. AF487494.. Corte-Real HB. Kalaydjieva L. Bosch E. 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