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Y-DNA haplogroups in populations of Europe

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File:Haplogroups europe.png
Y-haplogroups in Europe

Frequencies in some ethnic groups in Europe

The table below shows the human Y-chromosome DNA haplogroups, based on relevant studies, for various ethnic and other notable groups from Europe. The samples are taken from individuals identified with the ethnic and linguistic designations shown in the first two columns; the third column gives the Total Sample Size studied; and the other columns give the Percentage for each particular haplogroup. (Ethnic groups from the North Caucasus are considered in their own article.)

Population Language[1] n R1b R1a I  E1b1b J G N T Others Reference
Albanians IE (Albanian) 51 17.6 9.8 19.6 21.6 23.5 2.0 0.0 0.0 — Semino2000[2]
Albanians (Kosovar) IE (Albanian) 114 21.10 4.42 I1=5.31
I2a2=2.65 		47.37 J2=16.7 0 0 0 P[xQ,R1]=1.77 Pericic2005[3]
Albanians (Tirana) IE (Albanian) 30 13.3 13.3 16.7 23.3 20.0 3.3 — — — Bosch2006[4]
Albanians IE (Albanian) 55 18.2 9.1 I1=3.6
I2a=14.5
I2b=3.6 		25.5 J1=3.6
J2=20.0 		1.8 0.0 0.0 — Battaglia2008[5]
Albanians (Macedonia) IE (Albanian) 64 18.8 1.6 I1=4.5
I2a=12.5 		39.1 J1=6.3
J2=15.6 		1.6 0.0 0.0 — Battaglia2008[5]
Albanians IE (Albanian) 106 — — 23.6 — — — — — — Rootsi2004[6]
Arkhangelsk (Russia) Slavic, Uralic[citation needed] 28 0 17.9 50.0 3.6 0 0 28.6 0 0 Mirabal2009[7]
Andalusians IE (Italic) 29 65.5 0.0 — — — 0.0 0.0 6.9 L=3.4 Semino2000[2]
Andalusians IE (Italic) 103 — — 3.9 — — — — — — Rootsi2004[6]
Andalusians IE (Italic) 76 — — — 9.2 1.1 — — — — Semino2004[8]
Aromuns (Dukasi, Albania) IE (Italic) 39 2.6 2.6 17.9 17.9 48.7 10.3 0.0 0.0 — Bosch2006[4]
Aromuns (Andon Poci, Albania) IE (Italic) 19 36.8 0.0 42.1 15.8 5.3 0.0 0.0 0.0 — Bosch2006[4]
Aromuns (Kruševo, Macedonia) IE (Italic) 43 27.9 11.6 20.9 20.9 11.6 7.0 0.0 0.0 — Bosch2006[4]
Aromuns (Štip, Macedonia) IE (Italic) 65 23.1 21.5 16.9 18.5 20.0 0.0 0.0 0.0 — Bosch2006[4]
Aromuns (Romania) IE (Italic) 42 23.8 2.4 19.0 7.1 33.3 0.0 — — — Bosch2006[4]
Ashkenazi Jews IE (Germanic, West) 79 — 12.7 — 22.8 43.0 — — — — Nebel2001[9]
Ashkenazi Jews IE (Germanic, West) 442 — — 4.1 19.7 38.0 9.7 0.2 — Behar2004[10]
Austrians IE (Germanic, West) 219 32 14 — — — — — — —
Bashkirs (Perm) Altaic (Turkic) 43 86.0 9.3 0.0 0.0 0.0 2.3 2.3 0.0 — Lobov[11]
Basque (French, Spanish) Basque (Basque) 67 88.1 0.0 7.5 2.2 3.0 0.0 0.0 0.0 — Semino2000[2]
Bavarians IE (Germanic, West) 80 50.0 15.0 — 8.0 5.0 — 0.0 0.0 — Rosser2000[12]
Belgians IE (Germanic/Italic) 92 63.0 4.0 — 2.0 — — — — — Rosser2000[12]
Belarusians IE (Slavic, East) 41 0.0 39.0 10.0 — — 2.4 — — Rosser2000[12]
Belarusians IE (Slavic, East) 147 19.0 — — — — Rootsi2004[6]
Belarusians IE (Slavic, East) 68 4.4 45.6 25.0 4.4 — 1.5 8.8 — — Kharkov2005[13]
Belarusians IE (Slavic, East) 306 4.2 51.0 — 4.6 3.3 — 9.5 — — Behar2003[14]
Bearnais IE (Italic) 26 — — 7.7 — 3.8 — — — — Semino2004[8]
Bearnais IE (Italic) 43 — — — 3.7 — — — — — Rootsi2004[6]
Bosnians IE (Slavic, South) 69 1.4 24.6 42.0 10.1 0.0 — — — — Pericic2005[3]
British IE (Germanic, West) 32 68.8 9.4 — — — — — — — Helgason2000[15]
Bulgarians IE (Slavic, South) 127 11.0 17.3 27.5 19.7 18.1 1.6 — 0.8 — Karachanak2009[16]
Bulgarians IE (Slavic, South) 34 — 14.7 — — — — — — — Malaspina2003[17]
Bulgarians IE (Slavic, South) 116 — — — 20.7 — — — — — Cruciani2004[18]
Catalans IE (Italic) 24 79.2 0.0 4.2 4.2 4.2 8.0 — — — Rootsi2004d[19]
Catalans IE (Italic) 33 — — — 6.1 3.6 — — — — Semino2004[8]
Cantabrians (Pasiegos) IE (Italic) 56 — — — 42.9 — — — — — Cruciani2004[18]
Chuvashes Altaic (Turkic) 79 3.8 31.6 11.3 0 24.2 0 27.8 0 — Tambets2004[20]
Croats (mainland) IE (Slavic, South) 108 15.7 34.3 37.0 5.6 1.9 0.9 — — — Pericic2005[3]
Croat (mainland) IE (Slavic, South) 189 — — 38.1 — — — — — — Rootsi2004[6]
Cypriots IE (Greek) 45 9.0 2.0 — 27.0 — — — — — Rosser2000[12]
Czechs IE (Slavic, West) 257 — 34.2 18.3 5.8 4.7 5.1 1.6 — — Luca2007[21]
Czechs and Slovaks IE (Slavic, West) 45 35.6 26.7 — 2.2 — — — — — Semino2000[2]
Czechs and Slovaks IE (Slavic, West) 198 — — 13.6 — — — — — — Rootsi2004[6]
Danes IE (Germanic, North) 12 41.7 16.7 — — — — — — — Helgason2000[15]
Danes IE (Germanic, North) 194 — — 38.7 — — — — — — Rootsi2004[6]
Danes IE (Germanic, North) 35 — — — 2.9 — — — — — Cruciani2004[18]
Dutch IE (Germanic, West) 27 70.4 3.7 — — — — — — — Semino2000[2]
Dutch IE (Germanic, West) — — 26.7[6] 8.0[12] 0[8] — — — —
Dutch IE (Germanic, West) 410 50.2 3.3 32.9 2.9 5.1 4.1 0.2 — — Barjesteh2008[22]>
English (Central) IE (Germanic, West) 215 61.9 3.3 — — — — — — — Weale2002[23]
Estonians Uralic (Finnic) 207 9.0 — — 3.0 1.0 — 40.6 — — Rosser2000[12]
Estonians Uralic (Finnic) 118 — 37.3 — — — — — — — Laitinen2002[24]
Estonians Uralic (Finnic) 210 — — 18.6 — — — — — — Rootsi2004[6]
Finns Uralic (Finnic) 57 2.0 10.5 — 2.0 — — 63.2 — — Rosser2000[12]
Finns Uralic (Finnic) 38 0.0 7.9 28.9 — — — 63.2 — — [20]
French IE (Italic) 23 52.2 0 17.4 8.7 4.3 0 0 0 — Semino2000[2]
French IE (Italic) 40 — — — 8.0 — — — — — Rosser2000[12]
Frisians IE (Germanic, West) 94 56.0 7.0 29.0 2.0 6.0 — — — — Wilson2001[25]
Frisians IE (Germanic, West) 94 55.3 7.4 34.0 2.1 1.4 — — — — Weale2002[23]
Gagauz (Kongaz) Altaic (Turkic) 48 10.4 12.5 31.3 16.7 8.3 10.4 4.2 6.3 — Varzari2006[26]
Gagauz(Etulia) Altaic (Turkic) 41 14.6 26.8 24.4 9.8 7.3 17.1 0.0 0.0 — Varzari2006[26]
Germans IE (Germanic, West) 48 47.9 8.1 — — — — — — — Helgason2000[15]
Germans IE (Germanic, West) 16 50 6.2 37.5 6.2 0 0 0 0 — Semino2000[2]
Germans (Berlin) IE (Germanic, West) 103 23 22 32 10 2 — 2 — 9 Kayser2005[27]
Germans IE (Germanic) 1215 38.9 17.9 23.6 6.2 4.0 — 1.6 — 7.7 Kayser2005[27]
Greeks IE (Greek) 77 11.7 15.6 19.5 20.8 16.9 9.1 0.0 2.6 — Firasat2007[28]
Greeks IE (Greek) 118 22.8 8.3 — — — — — — — Helgason2000[15]
Greeks IE (Greek) 84/92 — — — 23.8 6.5 — — — — Semino2004[8]
Greeks IE (Greek) 171 13.5 11.1 15.8 31.6 19.9 4.7 — 1.8 — King2008[29]
Greeks (Crete) IE (Greek) 193 17.0 8.8 13.0 8.8 38.9 10.9 — 2.1 — King2008[29]
Greeks (Peloponnese) IE (Greek) 36 — — — 47 — — — — — Semino2004[8]
Greeks (Thrace) IE (Greek) 41 12.2 22.0 19.5 19.5 19.5 4.9 — — — Bosch2006[4]
Greeks (North) IE (Greek) 96 14.6 18.8 12.5 35.4 — 5.2 — 2.1 L=1 Zalloua2008[30]
Greeks (South) IE (Greek) 46 19.6 2.2 23.9 43.5 — 6.5 — 2.2 — Zalloua2008[30]
Gypsy (Macedonia) IE (Indic) 57 1.8 1.8 5.3 29.8 1.8 0 — — H=59.6 Pericic2005[3]
Herzegovinians IE (Slavic, South) 141 3.6 12.1 63.8 8.5 0.7 — — — — Pericic2005[3]
Hungarians Uralic (Ugric) 215 18.1 25.6 I1=7.91
I2a=16.74
I2b=2.79
I*=0.93 		6.1 J2=6.51
J*[xJ2]=3.72 		4.2 0.47 0.0 H=5.12
R2=0.47
R1*=1.40 		Völgyi2008[31]
Hungarians Uralic (Ugric) 45 13.3 60.0 11.1 8.9 2.2 2.2 0.0 0.0 L=2.2 Semino2000[2]
Hungarians Uralic (Ugric) — 56.6[5] 22.8[6] 9.4[8] 2.0[8] 8.0[32] 1.0 — —
Icelanders IE (Germanic, North) 181 41.4 23.8 34.2 — — — — — — Helgason2000[15]
Irish IE (Celtic) 222 81.5 0.5 — — — — — — — Helgason2000[15]
Irish IE (Celtic) 257 — — — 2.0 — — — — — Rosser2000[12]
Italians IE (Italic) 50 62.0 — 8.0 — — 10.0 — — — Rootsi2004d[19]
Italians IE (Italic) — 2.7[15] — 13.0[12] — — — — —
Italians (Calabria) IE (Italic) 32.4[2] — 5.4[6] 16.3[3] 24.6[8] — — — —
Italians (Apulia) IE (Italic) — — 2.6[6] 13.9[8] 31.4[8] — — — —
Italians (Sardinia) IE (Italic) 22.1[2] — 42.3[6] 5.0[8] 12.5[8] — — — —
Italians (Northern Sardinia) IE (Italic) 86 20.0 0.0 28.0 13.0 — 21.0 — 0.0 — Zalloua2008[30]
Italians (Southern Sardinia) IE (Italic) 187 19.0 1.0 35.0 11.0 — 14.0 — 0.0 — Zalloua2008[30]
Italians (North-central) IE (Italic) 62.0[2] — 0.5[6] 10.4[3] 26.9[8] — — — —
Italians (South) IE (Italic) 68 25.0 3.0 6.0 26.0 — 15.0 — 3.0 — Zalloua2008[30]
Italians (Sicily) IE (Italic) — — 8.8 27.3 23.8 — — — — Semino2004[8]
Italians (East Sicily) IE (Italic) 87 20.0 2.3 5.0 29.0 — 5.0 — 5.0 — Zalloua2008[30]
Italians (West Sicily) IE (Italic) 125 27.0 2.4 11.0 19.0 — 13.0 — 3.0 — Zalloua2008[30]
Komi Uralic (Finnic) 94 16.0 33.0 5.3 — — — 35.1 — — [20]
Komi (Izhemsky) Uralic (Finnic) 54 0.0 29.6 1.9 0.0 0.0 0.0 68.5 0.0 — Mirabal2009[7]
Komi (Priluzsky) Uralic (Finnic) 49 2.0 32.7 4.1 0.0 0.0 0.0 61.2 0.0 — Mirabal2009[7]
Kursk (Russia) Slavic 40 7.5 52.5 15.0 10.0 2.5 0 12.5 0 0 Mirabal2009[7]
Latvians IE (Baltic) 114 39.5 — 0.9 — — 42.1 — — Laitinen2002[24]
Latvians IE (Baltic) 34 15.0[12] 41[12] 7.0[12] — — — 32[12] — — Rosser 2000[12]
Lithuanians IE (Baltic) 38 5.0 — — — — — — — — [12]
Lithuanians IE (Baltic) 114 — 36.0 — 0.9 — — 43.0 — — Laitinen2002[24]
ethnic Macedonians IE (Slavic, South) 211 11.4 14.2 31.3 18.0 16.0 3.8 0.5 1.9 L=0.5 Noveski2010[33]
ethnic Macedonians IE (Slavic, South) 79 5.1 15.2 34.2 24.1 12.7 5.1 — — — Pericic2005[3]
Macedonians (Skopje) IE (Slavic, South) 52 13.5 13.5 28.8 23.1 11.5 3.8 — — — Bosch2006[4]
Maltese Afro-Asiatic (Semitic) 187 22.0 5.0 9.0 6.0 — 9.0 — 0.0 — Zalloua2008[30]
Mari Uralic (Finnic) 111 2.7 47.7 8.1 — — — 41.4 — — [20]
Mari Uralic (Finnic) 48 10.4 29.2 — 0.0 6.3 — 50.0 — — Rosser2000[12]
Moldavians (Karahasani) IE (Italic) 72 16.7 34.7 25.0 12.5 9.7 0.0 1.4 0.0 — Varzari2006[26]
Moldavians (Sofia) IE (Italic) 54 16.7 20.4 35.2 13.0 5.6 1.9 3.7 1.9 — Varzari2006[26]
Mordvins (Erzya) Uralic (Finnic) 46 — 39.1 — — — — — — — Malaspina2003[17]
Mordvins (Moksha) Uralic (Finnic) 46 — 21.7 — — — — — — — Malaspina2003[17]
Mordvins Uralic (Finnic) 83 13.3 26.5 19.3 — — — 19.3 — — [20]
Norwegians IE (Germanic, North) 25.9[15] 17.9[15] 40.3[6] — — — — — —
Norwegians IE (Germanic, North) 52 — 30.8 — 1.9 1.9 — 3.8 0.0 — Rosser2000[12]
Orcadians IE (Germanic, West) 71 66.0 19.7 — — — — — — — Wilson2001[25]
Poles IE (Slavic, West) 16.4[2] 56.4[2] 17.8[6] 4.0[8] 1.0[8] — — — —
Poles IE (Slavic, West) 93 13.4 55.9 16.1 — — — 3.2 — — [20]
Poles IE (Slavic, West) 913 11.6 57.0 17.3 4.5 2.5 — 3.7 — 3.3 Kayser2005[27]
Portuguese IE (Italic) 303 — — 5.3 — — — — — — Rootsi2004[6]
Portuguese (South) IE (Italic) 57 56.0 2.0 — 17.0 — — — — — Rosser2000[12]
Portuguese (North) IE (Italic) 328 62.0 0 — 11.0 — — — — — Rosser2000[12]
Romanians IE (Italic) 54 13.0 20.4 48.1 7.4 5.6 5.6 0.0 0.0 — Varzari2006[26]
Romanians IE (Italic) 361 — — 22.2 — — — — — — Rootsi2004[6]
Romanians (Constanţa) IE (Italic) 31 16.1 9.7 41.9 9.7 6.5 12.9 0.0 0.0 — Bosch2006[4]
Romanians (Ploieşti) IE (Italic) 36 8.3 5.6 38.9 16.7 19.4 8.3 0.0 0.0 — Bosch2006[4]
Russians IE (Slavic, East) 122 6.6 46.7 — 6.6 4.1 — 18.0 — — Rosser2000[12]
Russians IE (Slavic, East) 61 21.3 42.6 13.1 — — — 16.4 — — [20]
Russians (Northern) IE (Slavic, East) 380 6.1 33.4 14.5 0.3 1.8 1.3 41.3 0.0 — Balanovsky2008[34]
Russians (Central) IE (Slavic, East) 364 7.7 47.0 16.5 5.2 3.3 0.0 17.0 0.8 — Balanovsky2008[34]
Russians (Adygea) IE (Slavic, East) 78 — — 24.4 — — — — — — Rootsi2004[6]
Russians (Bashkortostan) IE (Slavic, East) 50 — — 6.0 — — — — — — Rootsi2004[6]
Russians (Belgorod region) IE (Slavic, East) 143 2.8 59.4 16.7 — — — — — — Balanovsky[34]
Russians (Cossacks) IE (Slavic, East) 97 — — 22.7 — — — — — — Rootsi2004[6]
Russians (Kostroma region) IE (Slavic, East) 53 — — 18.9 — — — — — — Rootsi2004[6]
Russians (North, Pinega) IE (Slavic, East) 127 — — 4.7 — — — — — — Rootsi2004[6]
Russians (Smolensk region) IE (Slavic, East) 120 — — 10.8 — — — — — — Rootsi2004[6]
Sami (Sweden) Uralic (Finnic) 38 7.9 15.8 31.6 0.0 0.0 0.0 44.7 0.0 — Karlsson2006[35]
Sami Uralic (Finnic) 127 3.9 11.0 — — — — 47.2 — — [20]
Sami Uralic (Finnic) — — 31.4 — — — — — — Rootsi2004[6]
Scots IE (Celtic) 178 — — 11.2 — — — — — — Rootsi2004[6]
Scots IE (Celtic) 61 77.1 6.6 — — — — — — — Helgason2000[15]
Sephardic Jews Afro-Asiatic (Semitic) 78 29.5 3.9 11.5 19.2 28.2 — — — — Nebel2001[9]
Serbs IE (Slavic, South) 179 4.5 14.5 48 17.3 5.6 2.2 3.3 — L=0.6 Mirabal,V.2010[36]
Serbs IE (Slavic, South) 113 10.6 15.9 36.3 21.2 8 — — — — Pericic2005[3]
Serbs IE (Slavic, South) 103 7.8 20.4 37.9 18.4 7.8 5.8 1.9 — — Regueiro2012[37]
Serbs (Bosnia) IE (Slavic, South) 81 6.2 13.6 40.7 22.2 9.9 1.2 6.2 0.0 — Battaglia2008[5]
Slovenians IE (Slavic, South) 75 21.3 38.7 30.7 2.7 4.0 2.7 0.0 0.0 — Battaglia2008[5]
Slovenian IE (Slavic, South) 70 — 37.1 — 7.1 5.7 — 0.0 0.0 — Rosser2000[12]
Slovenian IE (Slavic, South) 55 — — 38.2 — — — — — — Rootsi2004[6]
Spanish IE (Italic) 126 68.0 2.0 — 10.0 — — — — — Rosser2000[12]
Spanish (Ibiza) IE (Italic) 54 57.4 0.0 1.9 7.4 — 13.0 — 16.7 — Zalloua2008[30]
Spanish (Majorca) IE (Italic) 62 66.1 0.0 8.1 6.2 — 6.2 — 1.6 — Zalloua2008[30]
Spanish (Minorca) IE (Italic) 37 73.0 2.7 2.7 18.9 — 0.0 — 0.0 — Zalloua2008[30]
Spanish (South) IE (Italic) 162 65.0 2.0 6.0 9.0 — 4.0 — 0.0 — Zalloua2008[30]
Spanish (Valencia) IE (Italic) 73 64.0 3.0 10.0 11.0 — 1.0 — 1.0 — Zalloua2008[30]
Swedes (Northern) IE (Germanic, North) 48 22.9 18.8 — 2.1 2.1 — 8.3 — — Rosser2000[12]
Swedes IE (Germanic, North) 110 20.0 17.3 — — — — — — — Helgason2000[15]
Swedes IE (Germanic, North) 225 — — 40 — — — — — — Rootsi2004[6]
Swedes IE (Germanic, North) 160 13.1 24.4 37.5 1.3 0.0 — 14.4 — — Lappalainen2008[38]
Swiss IE (German/Italic) 144 — — 7.6 — — — — — — Rootsi2004[6]
Tatars Altaic (Turkic) 126 8.7 34.1 4.0 — — — 23.0 — — [20]
Turks (Istanbul) Altaic (Turkic) — — — 13.0 24.7 — — — — Semino2004[8]
Udmurt Uralic (Finnic) 87 2.3 10.3 1.1 — — — 85.1 — — [20]
Ukrainians IE (Slavic, East) 50 2.0 54.0 18 4 6 4.0 6.0 2.0 — Semino2000[2]
Ukrainians IE (Slavic, East) — — 21.9[6] 8.6[8] 7.3[8] — — — —
Ukrainians IE (Slavic, East) 53 18.9 41.5 24.5 — 9.4 0.0 5.7 0.0 — Varzari2006[26]
Welsh (Anglesey) IE (Celtic) 88 89.0 1.0 — 3.0 — — — — — Wilson2001[25]
Welsh (Anglesey) IE (Celtic) 196 — — 8.1 — — — — — — Rootsi2004[6]

Chronological development of haplogroups found in Europe

Listed here are notable groups from Europe and their chronological development based on relevant studies:[39][40][41][42][43][44][45][46][47][48][49][50][51][52][53][54][55][56][57][58][59][60][61][62][63][64][65][66][67][68][69][70][71][72][73][74][75][76][77][78][79][80][81][82][83][84][85][86][87][88][89][90][91][92][93][94][95][96][97][98][99][100][101][102][103][104][105][106][107][108][109][110][111][112][113][114][115][116][117][118][119][120][121][122][123][124][125][126][127][128][129][130][131]

Haplogroup Possible time of origin Possible place of origin Highest frequencies
K 40,000 years ago South Asia or West Asia
T 30,000 years ago West Asia
J 30,000 years ago Middle East
R 28,000 years ago Central Asia
E1b1b-M35 26,000 years ago East Africa
I 25,000 years ago Balkans
R1a1 21,000 years ago Southern Russia
R1b 20,000 years ago Around the Caspian Sea or Central Asia
E1b1b-M78 18,000 years ago Egypt/Libya
G 17,000 years ago Between India and the Caucasus
I2 17,000 years ago Balkans
J2 15,000 years ago Northern Mesopotamia
I2b 13,000 years ago Central Europe
N1c1 12,000 years ago Siberia or Mongolia 90% Yakuts
I2a 11,000 years ago Balkans
R1b1b2 10,000 years ago North or south of the Caucasus
J1 10,000 years ago Middle East
E1b1b-V13 8,500 years ago Middle East
I2b1 9,000 years ago Germany
I2a1 8,000 years ago Sardinia
I2a2 2,500 years ago Poland
E1b1b-M81 5,500 years ago Maghreb
I1 5,000 years ago Scandinavia
R1b-L21 4,000 years ago Central or Eastern Europe
R1b-S28 3,500 years ago around the Alps
R1b-S21 3,000 years ago Frisia or Central Europe
I2b1a < 3,000 years ago Britain

See also

References

  1. ^ IE = Indo-European
  2. ^ a b c d e f g h i j k l m n Ornella Semino et al 2000, The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective.
  3. ^ a b c d e f g h i Marijana Peričić et al 2005, High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations. Cite error: The named reference "Pericic2005" was defined multiple times with different content (see the help page).
  4. ^ a b c d e f g h i j Bosch, E.; Calafell, F.; González-Neira, A.; Flaiz, C; Mateu, E; Scheil, HG; Huckenbeck, W; Efremovska, L; Mikerezi, I (2006). "Paternal and maternal lineages in the Balkans show a homogeneous landscape over linguistic barriers, except for the isolated Aromuns". Annals of Human Genetics. 70 (Pt 4): 459–487. doi:10.1111/j.1469-1809.2005.00251.x. PMID 16759179.
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