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Particularly useful information is provided by the research of two uniparental markers within our genome, the <a href="/wiki/Y-chromosome" class="mw-redirect" title="Y-chromosome">Y-chromosome</a> (Y-DNA) and <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> (mtDNA), as well as <a href="/wiki/Autosomal_DNA" class="mw-redirect" title="Autosomal DNA">autosomal DNA</a>. The data from Y-DNA and autosomal DNA suggests that the <a href="/wiki/Croats" title="Croats">Croats</a> mostly are descendants of the <a href="/wiki/Slavs" title="Slavs">Slavs</a> of the medieval <a href="/wiki/Slavic_migrations_to_Southeastern_Europe" class="mw-redirect" title="Slavic migrations to Southeastern Europe">migration period</a>, according to mtDNA have genetic diversity which fits within a broader European maternal genetic landscape, and overall have a uniformity with other South Slavs from the territory of former <a href="/wiki/Yugoslavia" title="Yugoslavia">Yugoslavia</a>. There are many <a href="/wiki/Paleolithic" title="Paleolithic">Paleolithic</a> period sites located in the territory of Croatia, mostly ascribed to the <a href="/wiki/Mousterian" title="Mousterian">Mousterian</a> phase in the <a href="/wiki/Middle_Paleolithic" title="Middle Paleolithic">Middle Paleolithic</a> period. In the Neolithic period in <a href="/wiki/Southeast_Europe" title="Southeast Europe">Southeast Europe</a> were founded major cultures like <a href="/wiki/Vin%C4%8Da_culture" title="Vinča culture">Vinča</a>, <a href="/wiki/Varna_culture" title="Varna culture">Varna</a>, <a href="/wiki/Star%C4%8Devo_culture" title="Starčevo culture">Starčevo</a>. In the <a href="/wiki/Bronze_Age" title="Bronze Age">Bronze Age</a> happened symbiosis between <a href="/wiki/Proto-Indo-Europeans" title="Proto-Indo-Europeans">Proto-Indo-Europeans</a> of <a href="/wiki/Kurgan_hypothesis" title="Kurgan hypothesis">Kurgan culture</a> and autochthonous populations, leading to the formation among others also of Proto-<a href="/wiki/Illyrians" title="Illyrians">Illyrians</a>. They gradually mixed and were assimilated by the <a href="/wiki/Ancient_Rome" title="Ancient Rome">Romans</a>, <a href="/wiki/Celts" title="Celts">Celts</a>, <a href="/wiki/Ostrogoths" title="Ostrogoths">Ostrogoths</a>, and finally <a href="/wiki/Early_Slavs" title="Early Slavs">Early Slavs</a> since the late 6th century. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Y_chromosome_DNA">Y chromosome DNA</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=1" title="Edit section: Y chromosome DNA">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Ancient_Y-DNA_and_atDNA">Ancient Y-DNA and atDNA</h3>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=2" title="Edit section: Ancient Y-DNA and atDNA">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png/220px-PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png" decoding="async" width="220" height="302" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png/330px-PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/46/PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png/440px-PCA_and_Admixture_of_the_Croatian_Neolithic_Poto%C4%8Dani_samples.png 2x" data-file-width="750" data-file-height="1030" /></a><figcaption>PCA and Admixture of the Copper Age Potočani individuals with PCA of other Neolithic & Bronze & Iron Age samples compared to present-day Croats, per Novak, Olalde, Ringbauer et al. 2021.</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:292px;max-width:292px"><div class="trow"><div class="tsingle" style="width:144px;max-width:144px"><div class="thumbimage" style="height:130px;overflow:hidden"><a href="/wiki/File:West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1f/West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp/142px-West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp.png" decoding="async" width="142" height="130" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1f/West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp/213px-West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1f/West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp/284px-West_Eurasian_PCA_with_ancient_and_modern_Croatian_genomes.webp.png 2x" data-file-width="2006" data-file-height="1838" /></a></div></div><div class="tsingle" style="width:144px;max-width:144px"><div class="thumbimage" style="height:130px;overflow:hidden"><a href="/wiki/File:Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1f/Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp/142px-Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp.png" decoding="async" width="142" height="131" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1f/Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp/213px-Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1f/Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp/284px-Admixture_and_genetic_affinities_in_ancient_and_modern_Croatian_genomes.webp.png 2x" data-file-width="2007" data-file-height="1848" /></a></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">West Eurasian PCA's, Admixture and genetic affinities of ancient and modern Croatian genomes, per Freilich, Ringbauer, Los et al. 2021.</div></div></div></div> In the 2014 study, of the three successfully generated SNP profiles of Neolithic <a href="/wiki/Star%C4%8Devo_culture" title="Starčevo culture">Starčevo culture</a> samples from <a href="/wiki/Vinkovci" title="Vinkovci">Vinkovci</a>, two belonged to Y-DNA haplogroup G2a-P15 and one to I2a1-P37.2, which could indicate G2a as potential representatives of the spread of farming from the Near East to Europe, while I2a as Mesolithic substratum in Europe.<a href="#cite_note-FOOTNOTESzécsényi-Nagy_et_al.2015-1">[1]</a> In the 2018 study, 10 out of 17 samples from Croatia had a successful Y-DNA sequencing; two Croatia <a href="/wiki/Cardium_pottery" title="Cardium pottery">Cardial Neolithic</a> (6005–5786 BCE) samples from Zemunica Cave belonged to <a href="/wiki/Haplogroup_C-V20" title="Haplogroup C-V20">C1a2</a> and E1b1b1a1b1, Early-Neolithic Starčevo culture (5837–5659 BCE) from <a href="/wiki/Beli_Manastir" title="Beli Manastir">Beli Manastir</a>-Popova zemlja to C, Early-Neolithic Croatia <a href="/wiki/Cardium_pottery" title="Cardium pottery">Impressa culture</a> (5641–5560 BCE) from Kargadur to G2a2a1, two Middle-Neolithic <a href="/wiki/Sopot_culture" title="Sopot culture">Sopot culture</a> (5207–4546 BCE) samples from <a href="/wiki/Osijek" title="Osijek">Osijek</a> to G2a2a1 and J2a1, Late-Neolithic Sopot (4790–4558 BCE) from Beli Manastir-Popova zemlja to I, two <a href="/wiki/Vu%C4%8Dedol_culture" title="Vučedol culture">Vučedol culture</a> (2884–2582 BCE) samples from Beli Manastir-Popova zemlja and Vučedol Tell to R1b1a1a2a2 and G2a2a1a2a, and the Early-Middle Bronze Age (1631–1521 BCE) sample from Veliki Vanik belonged to J2b2a.<a href="#cite_note-FOOTNOTEMathieson_et_al.2018-2">[2]</a> In 2019 were autosomally analyzed three 5th century individuals with <a href="/wiki/Artificial_cranial_deformation" title="Artificial cranial deformation">artificial cranial deformation</a> from Osijek, probably of Hunnic or Germanic origin. According to <a href="/wiki/Principal_component_analysis" title="Principal component analysis">Principal component analysis</a> and Admixture methods, one of them had East/Northeast Asian, another European/Caucasian and third Near Middle Eastern/North African ancestry.<a href="#cite_note-FOOTNOTEFernandes_et_al.2019-3">[3]</a> In the 2021, was made a genome-wide analysis on 38 out of 41 individuals from the 6200 years old massacre at village <a href="/wiki/Velika,_Croatia" title="Velika, Croatia">Potočani</a> in Eastern Croatia.<a href="#cite_note-4">[4]</a><a href="#cite_note-5">[5]</a> They belonged to the Middle <a href="/wiki/Copper_Age" class="mw-redirect" title="Copper Age">Copper Age</a> <a href="/w/index.php?title=Lasinja_culture&action=edit&redlink=1" class="new" title="Lasinja culture (page does not exist)">Lasinja culture</a>, and autosomally 70% of them were not in close kinship implying "a community composed of many family groups". According to Admixture and PCA it revealed "that the analyzed individuals are slightly shifted from the Anatolia Neolithic cluster in the direction of Western European hunter-gatherers, similar to other Middle to Late Neolithic European farmers before the arrival of <a href="/wiki/Western_Steppe_Herders" title="Western Steppe Herders">[Western] steppe</a> ancestry", as on average had approximately 91% <a href="/wiki/Early_European_Farmers" title="Early European Farmers">Anatolian Neolithic Farmers</a> (ANF) and 9% <a href="/wiki/Western_Hunter-Gatherer" class="mw-redirect" title="Western Hunter-Gatherer">Western Hunter-Gatherer</a> (WHG)-related ancestry, being very distant from modern Croats. Among them were present "paternal lineages typical of Balkan Neolithic populations (eleven G2a2, two I2a1a(xI2a1a2), one I2a2, C and C1a2-V20)".<a href="#cite_note-FOOTNOTENovak,_Olalde,_Ringbauer_et_al.2021-6">[6]</a> In the same year were analyzed additional 19 Middle Neolithic of Sopot culture, 1 Copper Age and 1 Roman age sample from Beli Manastir-Popova zemlja as well as 7 Bronze Age samples of <a href="/w/index.php?title=Transdanubian_Encrusted_Pottery_culture&action=edit&redlink=1" class="new" title="Transdanubian Encrusted Pottery culture (page does not exist)">Transdanubian Encrusted Pottery culture</a> from <a href="/wiki/Jagodnjak" title="Jagodnjak">Jagodnjak</a>-Krčevine in Eastern Croatia. According to Admixture, Croatian Neolithic samples have 97.6-100% ANF-ancestry, Copper Age samples have 71% ANF-ancestry and 29% <a href="/wiki/Western_Steppe_Herders" title="Western Steppe Herders">Western Steppe Herders</a> (WSH)-ancestry, while Bronze Age samples 47% ANF, 20% WHG and 33% WSG-ancestry. The observed Y-DNA haplogroups in Neolithic/Copper period were two G2a2a-PF3147, one G2a2b2a1a1-PF3345, one C1a2b-Z38888, two I2a2a-M223 and one J-M304; in Bronze period were only four G2a2a1a2a2a1-Z31430 and one G2a2a1-PF3148. The one Roman period individual from 260 to 402 CE, "who may or may not be representative for the wider population in that time period", had autosomal genetic combination very similar to present-day Southeastern Europeans (including Croats), "adding an important data point to the substantial shift in ancestry that occurred in the region between the Bronze Age and today". He carried <a href="/wiki/Scythians#Archaeogenetics" title="Scythians">Scythian</a>-Eurasian Y-DNA haplogroup <a href="/wiki/Haplogroup_R1a#R1a1a1b2_(R-Z93)_(Asia)" title="Haplogroup R1a">R1a1a1b2a2-Z2124</a> > R1a1a1b2a2b1-F1345 (R-F1019* subclade<a href="#cite_note-7">[7]</a>).<a href="#cite_note-FOOTNOTEFreilich,_Ringbauer,_Los_et_al.2021-8">[8]</a> Between 2018 and 2022 studies analyzed many samples related to the <a href="/wiki/Cetina_culture" title="Cetina culture">Cetina culture</a>, Proto-<a href="/wiki/Illyrians" title="Illyrians">Illyrians</a> and specific Illyrian tribes (<a href="/wiki/Iapydes" class="mw-redirect" title="Iapydes">Iapydes</a> and <a href="/wiki/Liburni" class="mw-redirect" title="Liburni">Liburni</a>), finding that their paternal lineage almost exclusively belonged to the J2b-L283 haplogroup.<a href="#cite_note-FOOTNOTEMathieson_et_al.2018-2">[2]</a><a href="#cite_note-FOOTNOTEPatterson_et_al.2022-9">[9]</a><a href="#cite_note-FOOTNOTELazaridis_et_al.2021-10">[10]</a> A 2022 <a href="/wiki/Archaeogenetic" class="mw-redirect" title="Archaeogenetic">archaeogenetic</a> study published in <a href="/wiki/Science_(journal)" title="Science (journal)">Science</a> compared ancient, medieval, and modern Balkans population samples and found that the <a href="/wiki/Slavic_migrations_to_Southeastern_Europe" class="mw-redirect" title="Slavic migrations to Southeastern Europe">medieval Slavic migrations</a> "profoundly affected the region", resulting in the reduction of Anatolian Neolithic ancestry in Southeastern Europe. Pre-Slavic Balkan populations have the most of the Anatolian Neolithic component of ancestry, whereas present-day Slavs outside the Balkans have the least, "with present-day people from Southeastern Europe", including Croats, "intermediate between the two extremes", with Greeks and Albanians having the most while Croats and Hungarians the least.<a href="#cite_note-Lazaridis2022-11">[11]</a> A 2023 archaeogenetic study published in <a href="/wiki/Cell_(journal)" title="Cell (journal)">Cell</a> confirmed that the spread of Slavic language and identity was because of large movements of people of both males and females with specific Eastern European ancestry and that "more than half of the ancestry of most peoples in the Balkans today comes from the Slavic migrations, with around a third Slavic ancestry even in countries like Greece where no Slavic languages are spoken today".<a href="#cite_note-12">[12]</a><a href="#cite_note-Olalde2023-13">[13]</a> The pre-Slavic period population of Croatia and the region had Y-DNA haplogroups E-V13, R1b, J2b, J2a, G2a while with Slavic migration arrived R1a-Z282 and I2a-L621 among others.<a href="#cite_note-Olalde2023-13">[13]</a> <div class="mw-heading mw-heading3"><h3 id="Contemporary_Y-DNA">Contemporary Y-DNA</h3>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=3" title="Edit section: Contemporary Y-DNA">edit</a>]</div> <a href="/wiki/Genetics" title="Genetics">Genetically</a>, on the paternal <a href="/wiki/Y_chromosome" title="Y chromosome">Y chromosome</a> line and studies published between 2003 and 2008, a majority (>75%) of male Croats from Croatia belongs to one of the three major European Y-DNA <a href="/wiki/Haplogroup" title="Haplogroup">haplogroups</a> – <a href="/wiki/Haplogroup_I-M438" title="Haplogroup I-M438">I2</a> (32%<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTERootsi_et_al.2004-15">[15]</a><a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a>-34%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>), <a href="/wiki/Haplogroup_R1a_(Y-DNA)" class="mw-redirect" title="Haplogroup R1a (Y-DNA)">R1a</a> (27%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>-34%<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTERootsi_et_al.2004-15">[15]</a><a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a>) and <a href="/wiki/Haplogroup_R1b_(Y-DNA)" class="mw-redirect" title="Haplogroup R1b (Y-DNA)">R1b</a> (12%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>-15%<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTERootsi_et_al.2004-15">[15]</a><a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a>), while a minority (>25%) belongs to haplogroups <a href="/wiki/Haplogroup_E_(Y-DNA)" class="mw-redirect" title="Haplogroup E (Y-DNA)">E</a> (9%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>), <a href="/wiki/Haplogroup_I-M253" title="Haplogroup I-M253">I1</a> (5%<a href="#cite_note-FOOTNOTERootsi_et_al.2004-15">[15]</a>-9%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>), <a href="/wiki/Haplogroup_J_(Y-DNA)" title="Haplogroup J (Y-DNA)">J</a> (4%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>), <a href="/wiki/Haplogroup_N_(Y-DNA)" class="mw-redirect" title="Haplogroup N (Y-DNA)">N</a> (2%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>), and <a href="/wiki/Haplogroup_G_(Y-DNA)" class="mw-redirect" title="Haplogroup G (Y-DNA)">G</a> (1%<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a>). According to recent and more extensive studies published between 2012 and 2022, a majority (65%) of male Croats from Croatia belongs to haplogroups I2 (39%-40%) and R1a (22%-24%), while a minority (35%) belongs to haplogroups E (10%), R1b (6%-7%), J (6%-7%), I1 (5-8%), G (2%), <a href="/wiki/Haplogroup_Q-M242" title="Haplogroup Q-M242">Q</a> (0-1.93%), <a href="/wiki/Haplogroup_H_(Y-DNA)" title="Haplogroup H (Y-DNA)">H</a> (0-1.8%), <a href="/wiki/Haplogroup_T-M184" title="Haplogroup T-M184">T</a> (0.6%), N (0-0.6%) and <a href="/wiki/Haplogroup_L-M20" title="Haplogroup L-M20">L</a> (0.2%).<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg/220px-The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg" decoding="async" width="220" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/32/The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg/330px-The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/32/The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg/440px-The_approximate_frequency_and_variance_of_haplogroup_I-P37_clusters_in_Eastern_Europe.jpg 2x" data-file-width="2812" data-file-height="2404" /></a><figcaption>The approximate frequency and variance distribution of haplogroup I-P37 clusters, ancestral "Dnieper-Carpathian" (DYS448=20) and derived "Balkan" (DYS448=19: represented by a single SNP <a href="/wiki/Haplogroup_I-M438#I2a-L621" title="Haplogroup I-M438">I-PH908</a>), in Eastern Europe per O.M. Utevska (2017).</figcaption></figure> <a href="/wiki/Haplogroup_I_(Y-DNA)" class="mw-redirect" title="Haplogroup I (Y-DNA)">Haplogroup I</a> among Croats from Croatia is around 43.5%,<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> while in <a href="/wiki/%C5%A0okci" title="Šokci">Šokci</a>, a subgroup of Croats from eastern Croatia, is between 44% and 50% of the same haplogroup.<a href="#cite_note-Juric06-20">[20]</a><a href="#cite_note-Juric08-21">[21]</a> It is divided in two major subdivisions – subclade <a href="/wiki/Haplogroup_I2_(Y-DNA)" class="mw-redirect" title="Haplogroup I2 (Y-DNA)">I2</a> (37.7%), typical for the populations of the Balkans and Eastern Europe, and <a href="/wiki/Haplogroup_I1_(Y-DNA)" class="mw-redirect" title="Haplogroup I1 (Y-DNA)">I1</a> (5.8%), typical for the populations of Scandinavia.<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> From the I2 subclade, the most prevailing is I2a1a (I-P37.2) i.e. its subclade lineage I-M423 (18,400-14,000 YBP) > I-Y3104 (14,000-11,400 YBP) > I2a1a2b-L621 (11,400-6,500 YBP) > I-CTS10936 (6,500-5,600 YBP) > I-S19848 (5,600 YBP) > I-CTS4002 (5,600-5,100 YBP) > I-CTS10228 (5,100-3,400 YBP) > I-Y3120 (3,400-2,100 YBP) > I-S17250 (2,100-1,850 YBP) > <a href="/wiki/Haplogroup_I-M438#I2a-L621" title="Haplogroup I-M438">I-PH908</a> (1,850-1,700 YBP),<a href="#cite_note-YFull-22">[22]</a> which is typical of the South Slavic populations of <a href="/wiki/Southeastern_Europe" class="mw-redirect" title="Southeastern Europe">Southeastern Europe</a>, being highest in <a href="/wiki/Bosnia-Herzegovina" class="mw-redirect" title="Bosnia-Herzegovina">Bosnia-Herzegovina</a> (>50%).<a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a> In Croatia the highest frequency is observed in <a href="/wiki/Dalmatia" title="Dalmatia">Dalmatia</a>, peaking in cities of <a href="/wiki/Split,_Croatia" title="Split, Croatia">Split</a> (50.48%), <a href="/wiki/Dubrovnik" title="Dubrovnik">Dubrovnik</a> (53%) and <a href="/wiki/Zadar" title="Zadar">Zadar</a> (60%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> as well southern Adriatic islands of <a href="/wiki/Vis_(island)" title="Vis (island)">Vis</a> (44.6%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> <a href="/wiki/Bra%C4%8D" title="Brač">Brač</a> and <a href="/wiki/Kor%C4%8Dula" title="Korčula">Korčula</a> (~55%), and <a href="/wiki/Hvar" title="Hvar">Hvar</a> (52.88-65%).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> It is around average in the cities of <a href="/wiki/Pula" title="Pula">Pula</a> (31.31%) and <a href="/wiki/Osijek" title="Osijek">Osijek</a> (27.6-40.91%),<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> some southern <a href="/wiki/Mljet" title="Mljet">Mljet</a> (33.3%), <a href="/wiki/Lastovo" title="Lastovo">Lastovo</a> (33.3%) and middle Adriatic islands of <a href="/wiki/Pa%C5%A1man" title="Pašman">Pašman</a> (37.2%) and <a href="/wiki/Pag_(island)" title="Pag (island)">Pag</a> (36.4%).<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> The frequency is lowest in the town of <a href="/wiki/Vara%C5%BEdin" title="Varaždin">Varaždin</a> (18%) in northwestern Croatia,<a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> in the western mountainous <a href="/wiki/%C5%BDumberak_Mountains" title="Žumberak Mountains">Žumberak</a> region (18.2%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> and in the middle-northern islands of <a href="/wiki/Cres" title="Cres">Cres</a> (3%), <a href="/wiki/Dugi_otok" class="mw-redirect" title="Dugi otok">Dugi otok</a> (11.4%), <a href="/wiki/Ugljan" title="Ugljan">Ugljan</a> (24.6%) and <a href="/wiki/Krk" title="Krk">Krk</a> (27%).<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> The highest frequency is found in <a href="/wiki/Croats_of_Bosnia_and_Herzegovina" title="Croats of Bosnia and Herzegovina">Croats of Bosnia and Herzegovina</a> (63.8-73.3%).<a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a><a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> Based on 8 STR marker genetic distances closest are populations of near countries, but also depending on method, Ukraine, Belarus, Poland and Russia.<a href="#cite_note-FOOTNOTEŠarac_et_al.2016Supplementary_Table_9-24">[24]</a> The subclade's I-P37.2 very high frequency in the Western Balkans diminishes in all directions.<a href="#cite_note-FOOTNOTEPrimorac_et_al.2011-25">[25]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> The population with haplogroup I migrated to Europe from the Middle East, approximately 25,000-13,000 years ago. It represents the Paleolithic and Mesolithic <a href="/wiki/European_early_modern_humans" class="mw-redirect" title="European early modern humans">population of hunter gatherers</a> in Europe. However, in comparison to older research which argued a prehistoric autochthonous origin of the contemporary haplogroup I2 in Croatia and the Balkans,<a href="#cite_note-30">[nb 1]</a> Battaglia et al. (2009) already observed highest variance of the haplogroup in <a href="/wiki/Ukraine" title="Ukraine">Ukraine</a>, and Zupan et al. (2013) noted that it suggests it arrived with <a href="/wiki/Migration_Period#Second_wave" title="Migration Period">Slavic migration</a> from the homeland which was in present-day Ukraine.<a href="#cite_note-FOOTNOTEZupan_et_al.2013-31">[30]</a> The recent research by O.M. Utevska (2017), concluded that the haplogroup <a href="/wiki/Microsatellite" title="Microsatellite">STR</a> <a href="/wiki/Haplotype" title="Haplotype">haplotypes</a> have the highest diversity in Ukraine, with ancestral STR marker result "DYS448=20" comprising "<a href="/wiki/Dnieper" title="Dnieper">Dnieper</a>-<a href="/wiki/Carpathian" class="mw-redirect" title="Carpathian">Carpathian</a>" cluster, while younger derived result "DYS448=19" comprising the "Balkan cluster" which is predominant among the South Slavs.<a href="#cite_note-FOOTNOTEUtevska2017219–226,_302-32">[31]</a> This "Balkan cluster" also has the highest variance in Ukraine, which indicates that the very high frequency in the Western Balkan is because of a <a href="/wiki/Founder_effect" title="Founder effect">founder effect</a>.<a href="#cite_note-FOOTNOTEUtevska2017219–226,_302-32">[31]</a> Utevska calculated that the STR cluster divergence and its secondary expansion from the middle reaches of the Dnieper river or from <a href="/wiki/Eastern_Carpathians" class="mw-redirect" title="Eastern Carpathians">Eastern Carpathians</a> towards the Balkan peninsula happened approximately 2,860 ± 730 years ago, relating it to the times before Slavs, but much after the decline of the <a href="/wiki/Tripolye_culture" class="mw-redirect" title="Tripolye culture">Tripolye culture</a>.<a href="#cite_note-FOOTNOTEUtevska2017219–226,_302-32">[31]</a> However, STR-based calculations give overestimated dates,<sup id="cite_ref-FOOTNOTEŠarac_et_al.2016p._6:It_is_important_to_stress_that_the_proposed_old_age_of_the_I2a1b-M423_and_R1a1a1b1a*-M558_lineages_obtained_in_previous_studies_(Battaglia_et_al.,_2009;_Peričić_et_al.,_2005;_Rootsi_et_al.,_2004;_Underhill_et_al.,_2007,_2015)_has_been_based_on_STR_analysis_(8_and_10_loci,_respectively)_and_recent_studies_clearly_indicate_that_the_STR-based_age_calculations_tend_to_yield_overestimated_dates_(Batini_et_al.,_2015;_Hallast_et_al.,_2015;_Karmin_et_al.,_2015)._33-0" class="reference"><a href="#cite_note-FOOTNOTEŠarac_et_al.2016p._6:It_is_important_to_stress_that_the_proposed_old_age_of_the_I2a1b-M423_and_R1a1a1b1a*-M558_lineages_obtained_in_previous_studies_(Battaglia_et_al.,_2009;_Peričić_et_al.,_2005;_Rootsi_et_al.,_2004;_Underhill_et_al.,_2007,_2015)_has_been_based_on_STR_analysis_(8_and_10_loci,_respectively)_and_recent_studies_clearly_indicate_that_the_STR-based_age_calculations_tend_to_yield_overestimated_dates_(Batini_et_al.,_2015;_Hallast_et_al.,_2015;_Karmin_et_al.,_2015).-33">[32]</a><a href="#cite_note-34">[33]</a> and more specifically, the cluster is represented by a single SNP, I-PH908, known as I2a1a2b1a1a1c in <a href="/wiki/ISOGG" class="mw-redirect" title="ISOGG">ISOGG</a> phylogenetic tree (2019), and according to YFull YTree it formed and had TMRCA approximately 1,850-1,700 YBP.<a href="#cite_note-YFull-22">[22]</a> Although it is dominant among the modern Slavic peoples on the territory of the former Balkan provinces of the <a href="/wiki/Roman_Empire" title="Roman Empire">Roman Empire</a>, until now it was not found among the samples from the Roman period and is almost absent in contemporary population of <a href="/wiki/Italy" title="Italy">Italy</a>.<a href="#cite_note-Fóthi-35">[34]</a> According to Pamjav et al. (2019) and Fóthi et al. (2020), the distribution of ancestral subclades like of I-CTS10228 among contemporary carriers indicates a rapid expansion from Southeastern <a href="/wiki/Poland" title="Poland">Poland</a>, is mainly related to the Slavs and their medieval migration, and the "largest demographic explosion occurred in the Balkans".<a href="#cite_note-Fóthi-35">[34]</a><a href="#cite_note-HorolmaTibor2019-36">[35]</a> According to Olalde et al. (2023) it was confirmed that I2-L621 was absent before medieval period in the Balkans and arrived with the Slavs.<a href="#cite_note-Olalde2023-13">[13]</a> <a href="/wiki/Haplogroup_R1a#R-M417_(R1a1a1)" title="Haplogroup R1a">R1a1a1-M17</a> (22.1%-25.6%) is the second most prevailing haplogroup.<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> The haplogroup R-M17 in Croatia is mostly divided into two subclades, R-M558 which is predominant (19.2%), and <a href="/wiki/Haplogroup_R1a#R-M458_(R1a1a1b1a1)" title="Haplogroup R1a">R-M458</a> (4.9%), while <a href="/wiki/Haplogroup_R1a#R-Z282_(R1a1a1b1a)_(Eastern_Europe)" title="Haplogroup R1a">R-Z282</a> is rare (1.2%).<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> It has highest frequency in northern (29.1%) and central (23.6%) region, and almost the same frequency in eastern (18.6%), southern (19.1%), and western (20%) region of Croatia.<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> The highest local frequency of R1a1a1-M17 was observed in the Croats from Varaždin (38%) and Osijek (26-39%),<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> Žumberak (34.1),<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> and in the middle-northern islands of Dugi Otok (34.1%), Krk (37%), Pašman (38%) and Cres (56.6%),<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> being similar to the values of the other Slavs, like Slovenes, Czechs and Slovaks. The frequency is lower in Šokci from eastern Croatia (16%),<a href="#cite_note-Juric06-20">[20]</a><a href="#cite_note-Juric08-21">[21]</a> in the city of Dubrovnik (13.4%) and Split (19%) in Dalmatia, as well on the southern islands of Hvar (8-10.58%) and Vis (17%).<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> In Bosnian Croats, the frequency is similar to those of other South Slavs (12%).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEKovačević_et_al.2014-37">[36]</a> Considering subclades, the only outlier is island of Cres which had almost equal percentage of R1a-M558 (29.3%) and R1a-M458 (27.3%).<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> Based on 8 STR marker genetic distances closest are populations of near countries, but also depending on method, Belarus, Slovakia, Poland and Russia.<a href="#cite_note-FOOTNOTEŠarac_et_al.2016Supplementary_Table_9-24">[24]</a> The R-M558 subclade is more frequent among <a href="/wiki/East_Slavs" title="East Slavs">East Slavs</a> in Eastern Europe and Volga-Ural region, while R-M458 among <a href="/wiki/West_Slavs" title="West Slavs">West Slavs</a> in Central and Eastern Europe. Both are present in "informative frequencies in Balkan populations with known Slavonic heritage".<a href="#cite_note-38">[37]</a> R-M558 subclade CTS1211 was also found among Hungarian conquerors which indicates mixing and assimilation of the Slavs among the Hungarians.<a href="#cite_note-39">[38]</a> The highest frequency of the haplogroup <a href="/wiki/Haplogroup_R1b" title="Haplogroup R1b">Haplogroup R1b</a> (7.9%-9.1%), which in Croatia is divided into several subclades (mainly <a href="/wiki/Haplogroup_R-M269#R1b1a1a2a_(R-L23)" title="Haplogroup R-M269">R-L23</a> and its subclade <a href="/wiki/Haplogroup_R-M269#R-U152" title="Haplogroup R-M269">R-U152</a>), has in northern (10.9%) and central (11.8%) region of Croatia,<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> while locally in the Croats from the island of Krk (16.2%) and <a href="/wiki/Dugi_Otok" title="Dugi Otok">Dugi Otok</a> (25%),<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> and Žumberak (11.3%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> while in the southern islands, city of Dubrovnik (3.9%) and in Bosnian Croats it is almost absent (1-6%),<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> or like in Osijek it was not found.<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> These two haplogroups (R1a and R1b) are connected to <a href="/wiki/Proto-Indo-Europeans" title="Proto-Indo-Europeans">Proto-Indo-Europeans</a> migration from the Eurasian area some 5,000 years ago,<a href="#cite_note-FOOTNOTEPrimorac_et_al.2011-25">[25]</a> with R1a particularly to Slavic population's migration.<a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> Their frequency show north–south gradiation and an opposite frequency distribution to the haplogroup I-P37.2,<a href="#cite_note-FOOTNOTEPrimorac_et_al.2011-25">[25]</a> and the highest frequency is observed in the northern, central and eastern Croatia.<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:HgE1b1b1a2.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/HgE1b1b1a2.png/220px-HgE1b1b1a2.png" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/HgE1b1b1a2.png/330px-HgE1b1b1a2.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7a/HgE1b1b1a2.png/440px-HgE1b1b1a2.png 2x" data-file-width="1013" data-file-height="758" /></a><figcaption>The distribution of haplogroup <a href="/wiki/Haplogroup_E-V68#E-V13" title="Haplogroup E-V68">E-V13</a> in Europe.</figcaption></figure> From the haplogroup E (9.8%-10.6%) among Croats the most frequent is subclade <a href="/wiki/Haplogroup_E-V68#E-V13" title="Haplogroup E-V68">E1b1b1a1b-V13</a> (6.7%), while E1b1b1a3-M149 and <a href="/wiki/Haplogroup_E-M123" title="Haplogroup E-M123">E1b1b1c-M123</a> were also found in small numbers (1.1%).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> <a href="/wiki/Haplogroup_E-V68#E-V13" title="Haplogroup E-V68">E-V13</a> it's typical of the populations of south-eastern Europe, peaking among Kosovo Albanians (44%), and is also high among the Macedonians, Greeks, Romanians, Bulgarians and Serbs.<a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a> The highest frequency in Croatian mainland has been found in Varaždin (16%) and Žumberak (18.2%)<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> in central islands Dugi Otok (15.9%) and Ugljan (13.2%), as well southern islands <a href="/wiki/Vis_(island)" title="Vis (island)">Vis</a> (23.4%) and <a href="/wiki/Mljet" title="Mljet">Mljet</a> (15.4%).<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> In the northern islands of Cres (3%)<a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> and Krk (6.8%) was similar to other southern islands (3.7-4.3%).<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a> In Bosnian Croats the frequency was the same as among the Croats from Croatia (8.9%).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> Subclades of J1 are rare in Croatia, while J2 are higher in Croats from Croatia, peaking in Croats from Osijek (10.2%) and central islands Ugljan (10.2%) and Pašman (16.6%) as well the northern island of Krk (10.8%) and Cres (14.1%),<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> than in Bosnian Croats (both 1.1%).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> Subclade G2a-P15 both in Croatian and Bosnian Croats is found in low numbers (1.1%),<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> but peaks locally in the north-eastern town of Osijek (13.8%),<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> and the southern islands of Mljet (15.4%), Korčula (10.4%), Brač (6%) as well northern island Cres (7%).<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a> The haplogroup E and J are related to post-LGM, Neolithic migration of a population from Anatolia who brought with them domestication of wild animals and plants. Specifically, the haplogroup E's subclade probably arose locally in the Balkan not earlier than 8,000-10,000 years ago. These haplogroups show south–north gradiation.<a href="#cite_note-FOOTNOTEPrimorac_et_al.2011-25">[25]</a> The haplogroup G could have been present in Europe during the LGM or population with some of its subclades arrived with early farmers.<a href="#cite_note-FOOTNOTEPrimorac_et_al.2011-25">[25]</a> Haplogroup's <a href="/wiki/Haplogroup_N-M231" title="Haplogroup N-M231">N</a> subclades are rare in Croatia (0-2.2%).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> It is very frequent in the <a href="/wiki/Far_East" title="Far East">Far East</a>, like Siberia and China, while in Europe in Finns (60%) and in the Baltic countries (45%). Unusually for European populations, another central Asian-Siberian haplogroup <a href="/wiki/Haplogroup_Q-M242" title="Haplogroup Q-M242">Q</a> was found in unusually high frequencies due to founder effect in the islands of Hvar (7.69-14%), Lastovo (8.3%) and Korčula (6%).<a href="#cite_note-FOOTNOTEBarać_et_al.2003-14">[14]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2016-23">[23]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2022-19">[19]</a> It was also found in an individual from Herzegovina.<a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a> In later studies, a Croatian from Bosnia and Herzegovina was confirmed to belong to the subclade Q-L472>Z2902>B285>B29 found in Central-East Asia,<a href="#cite_note-40">[39]</a> while a Croatian from Croatia, another one from the island of Hvar, and a Croatian of Jewish origin in Poland belonged to the subclade Q-L245>Y2998>Y2209 which is mostly found in the Middle East and <a href="/wiki/Ashkenazi_Jews" title="Ashkenazi Jews">Ashkenazi Jews</a>.<a href="#cite_note-41">[40]</a> <div class="mw-heading mw-heading3"><h3 id="Abstract_and_data">Abstract and data</h3>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=4" title="Edit section: Abstract and data">edit</a>]</div> The region of modern-day Croatia was part of a wider Balkan region which may have served as one of several <a href="/wiki/Last_Glacial_Maximum_refugia" title="Last Glacial Maximum refugia">refugia</a> during the LGM, a source region for the recolonization of Europe during the post-glacial period and Holocene (10,000 years ago).<a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a><a href="#cite_note-FOOTNOTEPrimorac_et_al.2011-25">[25]</a> The eastern Adriatic coast was much further south.<a href="#cite_note-FOOTNOTERootsi_et_al.2004-15">[15]</a> The northern and the western parts of that sea were steppes and plains, while the modern Croatian islands (rich in Paleolithic archeological sites) were hills and mountains.<a href="#cite_note-FOOTNOTERootsi_et_al.2004-15">[15]</a><a href="#cite_note-FOOTNOTEBattaglia_et_al.2008-17">[17]</a> The region had a specific role in the structuring of European, and particularly among Slavic, paternal genetic heritage, characterized by the predominance of R1a and I, and scarcity of E lineages.<a href="#cite_note-FOOTNOTEPeričić_et_al.2005-16">[16]</a> The contemporary insular population's genetic diversity is characterized by strong isolation and <a href="/wiki/Endogamy" title="Endogamy">endogamy</a>.<a href="#cite_note-FOOTNOTEVitart_et_al.2006-42">[41]</a> In the table below is cited the most extensive study (2012) until now on the population in Croatia. It is a national reference <a href="/wiki/DNA_database" title="DNA database">DNA database</a> of 17 <a href="/wiki/Locus_(genetics)" title="Locus (genetics)">loci</a> system which acquired Y-<a href="/wiki/Microsatellite" title="Microsatellite">STR</a> <a href="/wiki/Haplotype" title="Haplotype">haplotypes</a> were predicted in estimated (over 90% probability) Y-<a href="/wiki/Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">SNP</a> <a href="/wiki/Haplogroup" title="Haplogroup">haplogroups</a>.<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> The sub-populations were divided in five regions which sub-populations showed strong similarity and homogeneity of paternal genetic contribution, all clustering together, with exception of sub-population from southern Croatia who showed a mild difference. In addition to high degree of overall homogeneity, there are gradient similarities to a central European cluster (Germanic, Slavic, Hungarian), and a southern European cluster (Bosnian, Serbian, Bulgarian, Macedonian, Romanian, Albanian, South Italian), going from north to south (and east to west).<a href="#cite_note-FOOTNOTEMršić_et_al.2012-18">[18]</a> Out of the neighboring countries and ethnic peoples are closest to them Bosnia and Herzegovina and <a href="/wiki/Bosniaks" title="Bosniaks">Bosniaks</a>.<a href="#cite_note-FOOTNOTEMršić_et_al.20127735-43">[42]</a><a href="#cite_note-44">[43]</a><a href="#cite_note-45">[44]</a> <table class="wikitable" style="text-align:center; font-size: 80%"> <tbody><tr> <th>Population</th> <th>Samples</th> <th>Source</th> <th><a href="/wiki/Haplogroup_I-M438#I2a-L621" title="Haplogroup I-M438">I2a</a>(xI2a1)</th> <th><a href="/wiki/Haplogroup_R1a" title="Haplogroup R1a">R1a</a></th> <th><a href="/wiki/Haplogroup_E-M215_(Y-DNA)" class="mw-redirect" title="Haplogroup E-M215 (Y-DNA)">E1b1b1-M35</a></th> <th><a href="/wiki/Haplogroup_R1b" title="Haplogroup R1b">R1b</a></th> <th><a href="/wiki/Haplogroup_I-M253" title="Haplogroup I-M253">I1</a></th> <th><a href="/wiki/Haplogroup_J-M172" title="Haplogroup J-M172">J2b</a></th> <th><a href="/wiki/Haplogroup_G-M201" title="Haplogroup G-M201">G2a</a></th> <th><a href="/wiki/Haplogroup_H_(Y-DNA)" title="Haplogroup H (Y-DNA)">H</a></th> <th><a href="/wiki/Haplogroup_J-M172" title="Haplogroup J-M172">J2a1h</a></th> <th><a href="/wiki/Haplogroup_J-M267" title="Haplogroup J-M267">J1</a></th> <th><a href="/wiki/Haplogroup_J-M172" title="Haplogroup J-M172">J2a1b</a></th> <th><a href="/wiki/Haplogroup_E-M2" title="Haplogroup E-M2">E1b1a1-M2</a></th> <th><a href="/wiki/Haplogroup_G-M285" title="Haplogroup G-M285">G1</a></th> <th><a href="/wiki/Haplogroup_G-M377" title="Haplogroup G-M377">G2c</a></th> <th><a href="/wiki/Haplogroup_I-M438#I-P37.2" title="Haplogroup I-M438">I2a1</a></th> <th><a href="/wiki/Haplogroup_I-M438#I-M223" title="Haplogroup I-M438">I2b1</a></th> <th><a href="/wiki/Haplogroup_I-M438#I-M223" title="Haplogroup I-M438">I2b(xI2b1)</a></th> <th><a href="/wiki/Haplogroup_J-M172" title="Haplogroup J-M172">J2a1-bh</a></th> <th><a href="/wiki/Haplogroup_L-M20" title="Haplogroup L-M20">L</a></th> <th><a href="/wiki/Haplogroup_N-M231" title="Haplogroup N-M231">N</a></th> <th><a href="/wiki/Haplogroup_Q-M242" title="Haplogroup Q-M242">Q</a></th> <th><a href="/wiki/Haplogroup_T-M184" title="Haplogroup T-M184">T</a> </th></tr> <tr> <td>Overall Croatia</td> <td>1,100</td> <td>Mršić et al. (2012)</td> <td>37.7% (415/1100)</td> <td>22.1% (243/1100)</td> <td>10.6% (117/1100)</td> <td>7.9% (87/1100)</td> <td>5.8% (64/1100)</td> <td>3.7% (41/1100)</td> <td>2.7% (30/1100)</td> <td>1.8% (20/1100)</td> <td>1.2% (13/1100)</td> <td>1.1% (12/1100)</td> <td>1% (11/1100)</td> <td>0.1%</td> <td>0</td> <td>0.1%</td> <td>0.1%</td> <td>0.9%</td> <td>0.3%</td> <td>0.3%</td> <td>0.2%</td> <td>0.6%</td> <td>0.7%</td> <td>0.6% </td></tr> <tr> <td>Central Croatia</td> <td>220</td> <td>Mršić et al. (2012)</td> <td>31.8% (70/220)</td> <td>23.6% (52/220)</td> <td>11.8% (26/220)</td> <td>10.4% (23/220)</td> <td>5% (11/220)</td> <td>5% (11/220)</td> <td>3.6% (8/220)</td> <td>1.3% (3/220)</td> <td>0.4% (1/220)</td> <td>2.2% (5/220)</td> <td>0.9% (2/220)</td> <td>0</td> <td>0</td> <td>0</td> <td>0.4%</td> <td>0.9%</td> <td>0.9%</td> <td>0</td> <td>0.4%</td> <td>0.9%</td> <td>0</td> <td>0 </td></tr> <tr> <td>North Croatia</td> <td>220</td> <td>Mršić et al. (2012)</td> <td>25.4% (56/220)</td> <td>29.1% (64/220)</td> <td>10.9% (24/220)</td> <td>10.4% (23/220)</td> <td>4.1% (9/220)</td> <td>5% (11/220)</td> <td>3.1% (7/220)</td> <td>5% (11/220)</td> <td>0.4% (1/220)</td> <td>0</td> <td>0.4% (1/220)</td> <td>0.4%</td> <td>0</td> <td>0</td> <td>0</td> <td>2.2%</td> <td>0.9%</td> <td>0</td> <td>0.4%</td> <td>0.4%</td> <td>0</td> <td>1.3% </td></tr> <tr> <td>East Croatia</td> <td>220</td> <td>Mršić et al. (2012)</td> <td>40% (88/220)</td> <td>18.6% (41/220)</td> <td>11.3% (25/220)</td> <td>8.2% (18/220)</td> <td>5.9% (13/220)</td> <td>2.7% (6/220)</td> <td>1.8% (4/220)</td> <td>0.9% (2/220)</td> <td>2.2% (5/220)</td> <td>2.7% (6/220)</td> <td>1.3% (3/220)</td> <td>0</td> <td>0</td> <td>0</td> <td>0</td> <td>0.4%</td> <td>0</td> <td>0.4%</td> <td>0</td> <td>0.9%</td> <td>1.8%</td> <td>0.4% </td></tr> <tr> <td>West Croatia</td> <td>220</td> <td>Mršić et al. (2012)</td> <td>36.8% (81/220)</td> <td>20% (44/220)</td> <td>12.7% (28/220)</td> <td>5.9% (13/220)</td> <td>8.6% (19/220)</td> <td>3.2% (7/220)</td> <td>3.2% (7/220)</td> <td>1.8% (4/220)</td> <td>1.8% (4/220)</td> <td>0.4% (1/220)</td> <td>1.8% (4/220)</td> <td>0</td> <td>0</td> <td>0.4%</td> <td>0</td> <td>0.4%</td> <td>0</td> <td>0.4%</td> <td>0</td> <td>0.9%</td> <td>0.4%</td> <td>0.9% </td></tr> <tr> <td>South Croatia</td> <td>220</td> <td>Mršić et al. (2012)</td> <td>54.5% (120/220)</td> <td>19.1% (42/220)</td> <td>6.3% (14/220)</td> <td>4.5% (10/220)</td> <td>5.4% (12/220)</td> <td>2.7% (6/220)</td> <td>1.8% (4/220)</td> <td>0.4% (1/220)</td> <td>0.9% (2/220)</td> <td>0</td> <td>0.4% (1/220)</td> <td>0</td> <td>0</td> <td>0</td> <td>0</td> <td>0.4%</td> <td>0</td> <td>0.9%</td> <td>0.4%</td> <td>0</td> <td>1.3%</td> <td>0.4% </td></tr> <tr> <td>Zagreb & Croatia</td> <td>239</td> <td>Purps et al. (2014)<a href="#cite_note-FOOTNOTEŠehović_et_al.2018-46">[45]</a></td> <td>36.1%</td> <td>23.8%</td> <td>6.4%</td> <td>13.9%</td> <td>3.9%</td> <td>2.1%</td> <td>n/a</td> <td>n/a</td> <td>1.9%</td> <td>0.9%</td> <td>n/a</td> <td>n/a</td> <td>0</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a </td></tr> <tr> <td>Croatia</td> <td>720</td> <td>Šarac et al. (2016)</td> <td>32.5%</td> <td>25.6%</td> <td>9.8%</td> <td>9.1%</td> <td>4.1%</td> <td>5.0%</td> <td>4.4%</td> <td>0.3%</td> <td>2.7%</td> <td>0.5%</td> <td>1.0%</td> <td>0.4%</td> <td>0</td> <td>0</td> <td>0.3%</td> <td>0.8%</td> <td>0.5%</td> <td>1.0%</td> <td>0</td> <td>0.6%</td> <td>0.9%</td> <td>1.2% </td></tr> <tr> <td>Croatia</td> <td>518</td> <td>Primorac et al. (2022)</td> <td>39%</td> <td>24.32%</td> <td>10.81%</td> <td>6.37%</td> <td>7.14%</td> <td>2.12%</td> <td>1.93%</td> <td>0</td> <td>3.47% (J2a)</td> <td>0.58%</td> <td>n/a</td> <td>0</td> <td>0.19%</td> <td>n/a</td> <td>1.35% (I2b)</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>0.19%</td> <td>0</td> <td>1.93%</td> <td>0.58% </td></tr> <tr> <td><a href="/wiki/West_Herzegovina_Canton" title="West Herzegovina Canton">West Herzegovina</a> (Croats)</td> <td>141</td> <td>Peričić et al. (2005)</td> <td>63.8%</td> <td>12%</td> <td>8.5%</td> <td>3.5%</td> <td>4.9%</td> <td>0.7%</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>0.7%</td> <td>n/a </td></tr> <tr> <td>Bosnia and Herzegovina (Croats)</td> <td>119</td> <td>Kovačević et al. (2014)<sup id="cite_ref-FOOTNOTEKovačević_et_al.2014:first_published_in_Marjanović_et_al._2005_with_90_samples,_the_same_samples_retested_in_Battaglia_et_al._2009,_retested_again_and_29_additional_samples_added_in_Kovačević_et_al._2014_47-0" class="reference"><a href="#cite_note-FOOTNOTEKovačević_et_al.2014:first_published_in_Marjanović_et_al._2005_with_90_samples,_the_same_samples_retested_in_Battaglia_et_al._2009,_retested_again_and_29_additional_samples_added_in_Kovačević_et_al._2014-47">[46]</a></td> <td>69.8%</td> <td>11.8%</td> <td>10.9%</td> <td>2.5%</td> <td>0</td> <td>0.8%</td> <td>0.8%</td> <td>n/a</td> <td>0.8%</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a</td> <td>n/a </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Mitochondrial_DNA">Mitochondrial DNA</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=5" title="Edit section: Mitochondrial DNA">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Ancient_mtDNA">Ancient mtDNA</h3>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=6" title="Edit section: Ancient mtDNA">edit</a>]</div> In the 2014 Y-DNA and mtDNA study, one Mesolithic sample dated 6080–6020 BCE from <a href="/wiki/Vela_Spila" title="Vela Spila">Vela Spila</a> near <a href="/wiki/Vela_Luka" title="Vela Luka">Vela Luka</a> on island <a href="/wiki/Kor%C4%8Dula" title="Korčula">Korčula</a> belonged to mtDNA haplogroup U5b2a5 common in hunter-gatherer communities, while other eleven Neolithic <a href="/wiki/Star%C4%8Devo_culture" title="Starčevo culture">Starčevo culture</a> samples dated circa 6000–5400 BCE from <a href="/wiki/Vinkovci" title="Vinkovci">Vinkovci</a> were assigned haplogroups J1c, K1a, T2b, HV0, K, V, V6, which reveal similar mtDNA diversity and shared ancestry in early farming populations from the <a href="/wiki/Pannonian_Basin" title="Pannonian Basin">Pannonian Basin</a> and the populations of the Central European <a href="/wiki/Linear_Pottery_culture" title="Linear Pottery culture">LBK</a>, accompanied by a reduction of the Mesolithic mtDNA substratum.<a href="#cite_note-FOOTNOTESzécsényi-Nagy_et_al.2015-1">[1]</a> Preliminary results from 2016 mtDNA study, which will approximately include 30 samples from Neolithic and 5 samples from Early to Late Bronze Age, on 5 ancient Croatian <a href="/wiki/Petrous_part_of_the_temporal_bone" title="Petrous part of the temporal bone">petrous bones</a> (3 Neolithic <a href="/wiki/Cardium_pottery" title="Cardium pottery">Cardial Impresso</a> from Zemunica Cave near <a href="/wiki/Bisko" title="Bisko">Bisko</a>, 1 Middle Neolithic <a href="/wiki/Danilo_culture" title="Danilo culture">Danilo culture</a> from Zidana Cave near Staničići Žumberački, 1 Mid/Late Bronze Age from Jazinka Cave near <a href="/wiki/Ne%C4%8Dven" title="Nečven">Nečven</a>) indicated mtDNA haplogroups K2 and K1b1a, H1e/H41, H1b for Neolithic samples similar to <a href="/wiki/Early_European_Farmers" title="Early European Farmers">Early European Farmers</a> (EEFs) and modern <a href="/wiki/Sardinians" class="mw-redirect" title="Sardinians">Sardinians</a> and Southern Europeans, while haplogroup HV or H4 for Bronze Age sample similar to modern day Croatian and Balkan population, but without clear evidence for connection with the Indo-European migration.<a href="#cite_note-FOOTNOTEFernandes_et_al.2016-48">[47]</a> The 2018 study which included 17 samples from Croatia; Mesolithic (7308–7027 BCE) from Vela Spila to U5b2b, three Croatia Cardial Neolithic (6005–5751 BCE) samples from Zemunica Cave to H1, K1b1a and N1a1, Early-Neolithic Starčevo (5837–5659 BCE) from Beli Manastir-Popova zemlja to U8b1b1, two Early-Neolithic Croatia Impressa (5670–5560 BCE) samples from Kargadur to H5a and H7c, two Middle-Neolithic Sopot (5207–4546 BCE) samples from Osijek to U5a1a2 and H10, two Late-Neolithic Sopot (4790–4536 BCE) samples from Beli Manastir-Popova zemlja to U5b2b and N1a1, Eneolithic (3710–3360 BCE) from Radovanci to J1c2, three <a href="/wiki/Vu%C4%8Dedol_culture" title="Vučedol culture">Vučedol</a> (3000–2582 BCE) samples from Beli Manastir-Popova zemlja and Vucedol Tell to T2e, T2c2 and U4a, Early-Middle Bronze Age (1631–1521 BCE) from Veliki Vanik to I1a1, and the Late Bronze Age (805–761 BCE) sample from Jazinka Cave belonged to HV0e.<a href="#cite_note-FOOTNOTEMathieson_et_al.2018-2">[2]</a> The 2021 study included 38 Neolithic (4200 BCE) samples from Potočani in Eastern Croatia and belonged to H, H4a1, H5b, H7, H13b1, H26, H42, HV, J1, K1a1, K1a3a, K1a4a1, K1b1b1, N1a1a1, N1a1a1a2, N1a1a1a3, T1a2, T2b, T2b23, T2f, U2, U5b1d1 and X2b, "suggesting that the Potočani victims belonged to a large community with a diverse pool of female lineages".<a href="#cite_note-FOOTNOTENovak,_Olalde,_Ringbauer_et_al.2021-6">[6]</a> In the same year study of 28 samples from Neolithic up to Roman times in Eastern Croatia found H, HV9, J2b1a5, K1a, K1a1, K1a1a, K1a4, K1a5, K1b1b1, K2a, K2b1, N1a1a1, N1a1a1a2, N1a1a1a3, T2b, T2b11, T2b21, T2b3, T2c1d1, T2f2, U2e1a1, U5a1g, U5b1b1a, U5b2b, U8b1a1.<a href="#cite_note-FOOTNOTEFreilich,_Ringbauer,_Los_et_al.2021-8">[8]</a> The 2011 mtDNA study on 27 early medieval skeletal remains in Naklice near <a href="/wiki/Omi%C5%A1" title="Omiš">Omiš</a> in Southern Dalmatia showed that 67% belonged to haplogroup H, 18% to J, 11% to U5, and 4% to HV.<a href="#cite_note-FOOTNOTELjubković_et_al.2011-49">[48]</a> The 2015 mtDNA study on medieval skeletal remains in <a href="/wiki/%C5%A0opot" title="Šopot">Šopot</a> (14th-15th century) and <a href="/wiki/Ostrovica,_Croatia" title="Ostrovica, Croatia">Ostrovica</a> (9th century) in Northern Dalmatia confirmed that profiles inherited by the maternal line differed neither between Ostrovica and Šopot site nor between medieval and modern populations, showing the same haplogroup prevalence (H, J, U) in both medieval and contemporary populations.<a href="#cite_note-FOOTNOTEBašić_et_al.2015-50">[49]</a> The 2014 study of a male skeleton found in <a href="/wiki/Split,_Croatia" title="Split, Croatia">Split</a> from Late Roman Period (dated 1760±80 YBP) showed that it belonged to haplogroup H.<a href="#cite_note-FOOTNOTESutlović_et_al.2014-51">[50]</a> <div class="mw-heading mw-heading3"><h3 id="Contemporary_mtDNA">Contemporary mtDNA</h3>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=7" title="Edit section: Contemporary mtDNA">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:World_map_of_prehistoric_human_migrations.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ab/World_map_of_prehistoric_human_migrations.jpg/220px-World_map_of_prehistoric_human_migrations.jpg" decoding="async" width="220" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ab/World_map_of_prehistoric_human_migrations.jpg/330px-World_map_of_prehistoric_human_migrations.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ab/World_map_of_prehistoric_human_migrations.jpg/440px-World_map_of_prehistoric_human_migrations.jpg 2x" data-file-width="889" data-file-height="635" /></a><figcaption>The mtDNA haplogroups hypothetical <a href="/wiki/Recent_African_origin_of_modern_humans" title="Recent African origin of modern humans">Out of Africa</a> migrations in thousand of years <a href="/wiki/Before_present" class="mw-redirect" title="Before present">before present</a>.</figcaption></figure> Genetically, on the maternal <a href="/wiki/Human_mitochondrial_DNA_haplogroup" title="Human mitochondrial DNA haplogroup">mitochondrial DNA</a> line, a majority (>65%) of Croats from Croatia (mainland and coast) belong to three of the eleven major European mtDNA haplogroups – <a href="/wiki/Haplogroup_H_(mtDNA)" title="Haplogroup H (mtDNA)">H</a> (25.5-45%), <a href="/wiki/Haplogroup_U_(mtDNA)" class="mw-redirect" title="Haplogroup U (mtDNA)">U</a> (18.85-22%), <a href="/wiki/Haplogroup_J_(mtDNA)" title="Haplogroup J (mtDNA)">J</a> (7.5-11%), while a large minority (>35%) belongs to many other smaller haplogroups.<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a><a href="#cite_note-FOOTNOTEBarbarić_et_al.2020-53">[52]</a> In all the studies, haplogroup H is the most frequent (45% in 2014) maternal haplogroup in Croatian mainland (continental) and coast respectively, but in most recent 2020 study is at lower frequencies of 25.5% due to nomenclature differences primarily of R/R0 (7.5%) lineages.<a href="#cite_note-FOOTNOTEBarbarić_et_al.2020-53">[52]</a> The highest frequency in Croatia observed in population of island <a href="/wiki/Kor%C4%8Dula" title="Korčula">Korčula</a> (60.2%), <a href="/wiki/Susak" title="Susak">Susak</a> (66%) and <a href="/wiki/Mljet" title="Mljet">Mljet</a> (73.5%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2017-54">[53]</a> while lowest frequency in islands <a href="/wiki/Cres" title="Cres">Cres</a> (27.7%),<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> and <a href="/wiki/Hvar" title="Hvar">Hvar</a> (27.8%).<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> It is the dominant European haplogroup.<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> The elevated frequency of subhaplogroup H1b in Mljet (30.9%), otherwise rare in other studies, is a typical example of a founder effect – migration from the nearest coastal region and micro-evolutionary expansion in the island.<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> Haplogroup U (18.85% in 2014,<a href="#cite_note-FOOTNOTEŠarac_et_al.2014-57">[56]</a> 22% in 2020<a href="#cite_note-FOOTNOTEBarbarić_et_al.2020-53">[52]</a>) is mostly represented by its subclade <a href="/wiki/Haplogroup_U_(mtDNA)#Haplogroup_U5" class="mw-redirect" title="Haplogroup U (mtDNA)">U5</a> which is the second most frequent haplogroup, with 11.6% in the mainland and 10.4% in the coast, with similar frequencies in the islands of <a href="/wiki/Bra%C4%8D" title="Brač">Brač</a> (8.6%), <a href="/wiki/Krk" title="Krk">Krk</a> (11.3%), and Hvar (14.8%), while lowest in Korčula (2%).<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> Overall the haplogroup U, including its subclades like U5, is the most frequent in the city of <a href="/wiki/Dubrovnik" title="Dubrovnik">Dubrovnik</a> (19.9%) and islands <a href="/wiki/Lastovo" title="Lastovo">Lastovo</a> (23.5%) and Cres (29.4%).<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> It is the oldest European haplogroup (coalescence age circa 50,000 YBP) and its subclade U5 makes the majority of the haplogroup diversity in Europe. The high frequency of <a href="/wiki/Haplogroup_U_(mtDNA)#Haplogroup_U4" class="mw-redirect" title="Haplogroup U (mtDNA)">U4</a> in Lastovo (11.8%) indicates founder effect.<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> Haplogroup J is the third most frequent haplogroup (7.5% in 2020<a href="#cite_note-FOOTNOTEBarbarić_et_al.2020-53">[52]</a>), with 11.9% in the mainland but only 3.1% in the coast, however the islands had higher frequencies than the coastal population Korčula (6.1%), Brač (8.6%), Krk (9%), Hvar (9.3%),<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> peak in <a href="/wiki/%C5%BDumberak_Mountains" title="Žumberak Mountains">Žumberak</a> (12.34%)<a href="#cite_note-FOOTNOTEŠarac_et_al.2014-57">[56]</a> and Lastovo (19.6%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> while in Cres is almost totally absent.<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> Haplogroup <a href="/wiki/Haplogroup_T_(mtDNA)" title="Haplogroup T (mtDNA)">T</a> is third or fourth most frequent haplogroup (10% in 2020<a href="#cite_note-FOOTNOTEBarbarić_et_al.2020-53">[52]</a>). Its subclade T2 has similar frequency of 3.1-5.8% in both the coastal and mainland as well insular population, with exceptional peak in island Hvar (12%),<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a><a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> and Susak (33%),<a href="#cite_note-FOOTNOTEŠarac_et_al.2017-54">[53]</a> however the overall haplogroup T has lower frequency in Mljet (1.5%), Lastovo (3.9%) and Dubrovnik (2.3).<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> Haplogroup <a href="/wiki/Haplogroup_K_(mtDNA)" title="Haplogroup K (mtDNA)">K</a> (7.5% in 2020<a href="#cite_note-FOOTNOTEBarbarić_et_al.2020-53">[52]</a>) has average frequency of 3.6% in the mainland and 6.3% in the coast, it is absent in Lastovo<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> and it has lowest frequency in the islands Cres (3.36%)<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> and Hvar (3.7%), while highest in the island Brač (9.5%).<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> Haplogroup <a href="/wiki/Haplogroup_V_(mtDNA)" title="Haplogroup V (mtDNA)">V</a> (under <a href="/wiki/Haplogroup_HV_(mtDNA)" class="mw-redirect" title="Haplogroup HV (mtDNA)">HV</a> > HV0) is a younger sister clade of haplogroup H,<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> and has almost the same minimum and maximum frequency (4-5.7%) in both continental and insular populations, with exception in Korčula (8.2%),<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> as well lower frequency in Mljet (1.5%), Lastovo (2%) and Dubrovnik (3.3%).<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> Haplogroup <a href="/wiki/Haplogroup_W_(mtDNA)" class="mw-redirect" title="Haplogroup W (mtDNA)">W</a> frequency in the mainland and coastal population is between 2.2 and 4.2%, while between insular populations 1.9-3.1%, with exception in Krk (7.5%),<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> and Cres (12.6%).<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> In islands Mljet and Lastovo is between 4.4 and 5.9%, while in Dubrovnik is almost absent (0.6%).<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> Other mtDNA haplogroup with notable local peaks are: HV subclades with low frequencies in the mainland and coast (0.4-2.1%) but average (4.1-4.6) in islands, and high in Dubrovnik (7.7%)<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> and Brač (10.5%).<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> Haplogroup <a href="/wiki/Haplogroup_N_(mtDNA)" title="Haplogroup N (mtDNA)">N1a</a> in Cres (9.24%) is the northernmost finding till now of this branch in Europe, and haplotypes indicate a relatively recent founder effect.<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> It is a characteristic haplogroup of the early farmers.<a href="#cite_note-FOOTNOTESzécsényi-Nagy_et_al.2015-1">[1]</a> Haplogroup <a href="/wiki/Haplogroup_F_(mtDNA)" title="Haplogroup F (mtDNA)">F</a> which is almost absent (only 0.2-0.4% in the mainland), but peaks at 8.3% in Hvar.<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a><a href="#cite_note-FOOTNOTEŠarac_et_al.2014-57">[56]</a> Haplogroup <a href="/wiki/Haplogroup_I_(mtDNA)" title="Haplogroup I (mtDNA)">I</a> in Krk (11.3%), which subhaplogroups separated around the LGM.<a href="#cite_note-FOOTNOTEHavaš_Auguštin_et_al.2012-58">[57]</a> Haplogroup <a href="/wiki/Haplogroup_X_(mtDNA)" title="Haplogroup X (mtDNA)">X</a> ranges 0.63-3.17%, mainly belonging to subclade X2 > X2b,<a href="#cite_note-FOOTNOTEŠarac_et_al.2014-57">[56]</a> and recent research of Cres and Rab possibly found a "new, island-specific" X3 lineage which "formed within the Croatian population".<a href="#cite_note-FOOTNOTEHavaš_Auguštin_et_al.2022-59">[58]</a> <div class="mw-heading mw-heading3"><h3 id="Abstract_and_data_2">Abstract and data</h3>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=8" title="Edit section: Abstract and data">edit</a>]</div> For decades the Croatian insular populations have been studied because of their isolation which can trace micro-evolutionary processes and understand evolutionary forces, like genetic drift (specific genetic expression), founder effect and population bottlenecks (reduction of population size) which shaped the contemporary population. The results until now indicate that the genetic flow and influx of women to the islands was limited.<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> A moderate genetic isolate can also be considered for the continental population of mountainous region <a href="/wiki/%C5%BDumberak_Mountains" title="Žumberak Mountains">Žumberak</a> because they had a loose affinity with <a href="/wiki/Uskoks" title="Uskoks">Uskoks</a>'s proposed region of origin (Bosnians, Herzegovinians, southern Croatians) or to their current closest neighbors (northern Croatians and Slovenians).<a href="#cite_note-FOOTNOTEŠarac_et_al.2014-57">[56]</a> On the example of population of the island of Krk, the high-resolution mtDNA analysis showed evidence that settlements <a href="/wiki/Omi%C5%A1alj" title="Omišalj">Omišalj</a>, <a href="/wiki/Vrbnik" title="Vrbnik">Vrbnik</a>, and <a href="/wiki/Dobrinj" title="Dobrinj">Dobrinj</a> are related in a joint cluster of early Slavic settlements, while <a href="/wiki/Poljica,_Krk" title="Poljica, Krk">Poljica</a> and <a href="/wiki/Duba%C5%A1nica" title="Dubašnica">Dubašnica</a> regions a separate cluster founded by Slavic and <a href="/wiki/Vlachs_of_Croatia" class="mw-redirect" title="Vlachs of Croatia">Vlachs</a> or <a href="/wiki/Morlachs" title="Morlachs">Morlachs</a> migrants from the Velebit hinterland who arrived in the 15th century.<a href="#cite_note-FOOTNOTEHavaš_Auguštin_et_al.2012-58">[57]</a> On the example of population of the island of Mljet can be perceived demographic and historical events like the island's use for quarantine station, while along Vis and Lastovo consanguinity practice and inbreeding due to lack of genetic diversity, being suitable for genetic-epidemiological research.<a href="#cite_note-FOOTNOTEŠarac_et_al.2012-52">[51]</a> In the 2004 mtDNA analysis, one cluster was formed by populations from islands Hvar, Krk and Brač, and second cluster included Croatian mainland and Croatian coast, while the island of Korčula was distinguished due to exceptionally high frequency of haplogroup H.<a href="#cite_note-FOOTNOTECvjetan_et_al.2004-56">[55]</a> In the 2009 mtDNA interpopulation PCA analysis of sub-haplogroups, insular populations from Krk, <a href="/wiki/Ugljan" title="Ugljan">Ugljan</a>, Korčula, Brač, Hvar were clustered together implying to have close maternal lineages, with Vis close to them, but <a href="/wiki/Rab_(G-protein)" title="Rab (G-protein)">Rab</a> (U4, H6, J1c) and especially Cres (prevalence U2, W, N1a) had separate outlying positions from both the cluster and each other, and confirmed "that genetic drift, especially founder effect, has played significant role in shaping genetic composition of the isolated population of the island of Cres".<a href="#cite_note-FOOTNOTEJeran_et_al.2009-55">[54]</a> In 2017 study the greatest outlier due to isolation, endogamy and lowest reported gene diversity index was Susak.<a href="#cite_note-FOOTNOTEŠarac_et_al.2017-54">[53]</a> In the 2014 mtDNA PCA analysis, the populations from eastern and southern Croatia clustered together with Bosnia and Herzegovina, while western and northern Croatia with Slovenia. As Slovenian population does not form Southeast Europe cluster it is considered a possible input from different migration waves of Slavs in the <a href="/wiki/Middle_Ages" title="Middle Ages">Middle Ages</a>.<a href="#cite_note-FOOTNOTEŠarac_et_al.2014-57">[56]</a> <table class="wikitable" style="text-align:center; font-size: 84%"> <tbody><tr> <th>Population</th> <th>Samples</th> <th>Source</th> <th><a href="/wiki/Haplogroup_H_(mtDNA)" title="Haplogroup H (mtDNA)">H</a></th> <th><a href="/wiki/Haplogroup_HV_(mtDNA)" class="mw-redirect" title="Haplogroup HV (mtDNA)">HV</a></th> <th><a href="/wiki/Haplogroup_J_(mtDNA)" title="Haplogroup J (mtDNA)">J</a></th> <th><a href="/wiki/Haplogroup_T_(mtDNA)" title="Haplogroup T (mtDNA)">T</a></th> <th><a href="/wiki/Haplogroup_K_(mtDNA)" title="Haplogroup K (mtDNA)">K</a></th> <th><a href="/wiki/Haplogroup_U_(mtDNA)" class="mw-redirect" title="Haplogroup U (mtDNA)">U</a>*</th> <th>U1</th> <th>U2</th> <th>U3</th> <th>U4</th> <th>U5</th> <th>U6</th> <th>U7</th> <th>U8</th> <th><a href="/wiki/Haplogroup_R_(mtDNA)" title="Haplogroup R (mtDNA)">R</a></th> <th><a href="/wiki/Haplogroup_N_(mtDNA)" title="Haplogroup N (mtDNA)">N</a></th> <th><a href="/wiki/Haplogroup_I_(mtDNA)" title="Haplogroup I (mtDNA)">I</a></th> <th><a href="/wiki/Haplogroup_W_(mtDNA)" class="mw-redirect" title="Haplogroup W (mtDNA)">W</a></th> <th><a href="/wiki/Haplogroup_X_(mtDNA)" title="Haplogroup X (mtDNA)">X</a></th> <th>Other </th></tr> <tr> <td>Croatia</td> <td>488</td> <td>Šarac et al. (2014)</td> <td>45.29</td> <td>4.07</td> <td>9.83</td> <td>5.98</td> <td>4.30</td> <td>0</td> <td>1.23</td> <td>2.66</td> <td>1.43</td> <td>2.66</td> <td>10.06</td> <td>0.20</td> <td>0.41</td> <td>0.20</td> <td>0</td> <td>0.82</td> <td>2.61</td> <td>1.84</td> <td>1.84</td> <td>4.29 </td></tr> <tr> <td>East Croatia</td> <td>61</td> <td>Šarac et al. (2014)</td> <td>49.18</td> <td>11.48</td> <td>9.84</td> <td>3.28</td> <td>4.92</td> <td>0</td> <td>0</td> <td>0</td> <td>1.64</td> <td>1.64</td> <td>9.84</td> <td>0</td> <td>0</td> <td>0</td> <td>0</td> <td>1.64</td> <td>0</td> <td>4.92</td> <td>1.64</td> <td>0 </td></tr> <tr> <td>North Croatia</td> <td>155</td> <td>Šarac et al. (2014)</td> <td>41.77</td> <td>5.06</td> <td>14.56</td> <td>10.76</td> <td>3.16</td> <td>0</td> <td>0.63</td> <td>1.90</td> <td>0.63</td> <td>3.16</td> <td>11.39</td> <td>0.63</td> <td>0</td> <td>0.63</td> <td>0</td> <td>0</td> <td>1.27</td> <td>2.53</td> <td>0.63</td> <td>0.63 </td></tr> <tr> <td>West Croatia</td> <td>209</td> <td>Šarac et al. (2014)</td> <td>46.41</td> <td>11.48</td> <td>6.70</td> <td>2.39</td> <td>5.74</td> <td>0</td> <td>2.39</td> <td>4.31</td> <td>1.91</td> <td>2.39</td> <td>6.22</td> <td>0</td> <td>0.96</td> <td>0</td> <td>0</td> <td>0.96</td> <td>4.78</td> <td>0</td> <td>2.39</td> <td>0 </td></tr> <tr> <td>South Croatia</td> <td>63</td> <td>Šarac et al. (2014)</td> <td>49.21</td> <td>3.17</td> <td>7.94</td> <td>9.52</td> <td>3.17</td> <td>0</td> <td>0</td> <td>1.59</td> <td>1.59</td> <td>3.17</td> <td>12.70</td> <td>0</td> <td>0</td> <td>0</td> <td>0</td> <td>1.59</td> <td>1.59</td> <td>1.59</td> <td>3.17</td> <td>0 </td></tr> <tr> <td>Croatia</td> <td>200</td> <td>Barbarić et al. (2020)</td> <td>25.5</td> <td>11.5</td> <td>7.5</td> <td>10</td> <td>7.5</td> <td>0</td> <td>2</td> <td>4</td> <td>2.5</td> <td>2.5</td> <td>10</td> <td>0</td> <td>1</td> <td>0</td> <td>7.5</td> <td>2</td> <td>3</td> <td>1</td> <td>2</td> <td>0.5 </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Contemporary_autosomal_DNA">Contemporary autosomal DNA</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=9" title="Edit section: Contemporary autosomal DNA">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Genetic_PCA_of_European_countries_2022.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/Genetic_PCA_of_European_countries_2022.jpg/220px-Genetic_PCA_of_European_countries_2022.jpg" decoding="async" width="220" height="167" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/Genetic_PCA_of_European_countries_2022.jpg/330px-Genetic_PCA_of_European_countries_2022.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/55/Genetic_PCA_of_European_countries_2022.jpg/440px-Genetic_PCA_of_European_countries_2022.jpg 2x" data-file-width="4189" data-file-height="3184" /></a><figcaption>The median plot of Croats from Croatia on a PCA including European countries, per Gilbert et al. 2022.<a href="#cite_note-FOOTNOTEGilbert_et_al.2022-60">[59]</a></figcaption></figure> According to 2013 <a href="/wiki/Autosome" title="Autosome">autosomal</a> <a href="/wiki/Identity_by_descent" title="Identity by descent">IBD</a> survey "of recent genealogical ancestry over the past 3,000 years at a continental scale", the speakers of Serbo-Croatian language share a very high number of common ancestors dating to the <a href="/wiki/Migration_period" class="mw-redirect" title="Migration period">migration period</a> approximately 1,500 years ago with Poland and Romania-Bulgaria cluster among others in Eastern Europe. It is concluded to be caused by the <a href="/wiki/Huns" title="Huns">Hunnic</a> and Slavic expansion, which was a "relatively small population that expanded over a large geographic area", particularly "the expansion of the Slavic populations into regions of low population density beginning in the sixth century" and that it is "highly coincident with the modern distribution of Slavic languages".<a href="#cite_note-FOOTNOTERalph_et_al.2013-61">[60]</a> According to Kushniarevich et al., the Hellenthal et al. 2014 IBD analysis also found "multi-directional admixture events among East Europeans (both Slavic and non-Slavic), dated to around 1,000–1,600 YBP" which coincides with "the proposed time-frame for the Slavic expansion".<a href="#cite_note-FOOTNOTEKushniarevich_et_al.2015-62">[61]</a> The 2015 IBD analysis found that the South Slavs have lower proximity to Greeks than with East and West Slavs, and that there's an "even patterns of IBD sharing among East-West Slavs–'inter-Slavic' populations (<a href="/wiki/Hungarians" title="Hungarians">Hungarians</a>, <a href="/wiki/Romanians" title="Romanians">Romanians</a> and <a href="/wiki/Gagauz_people" title="Gagauz people">Gagauz</a>)–and South Slavs, i.e. across an area of assumed historic movements of people including Slavs". The slight peak of shared IBD segments between South and East-West Slavs suggests a shared "Slavonic-time ancestry".<a href="#cite_note-FOOTNOTEKushniarevich_et_al.2015-62">[61]</a> The 2014 IBD analysis comparison of Western Balkan and Middle Eastern populations also found negligible gene flow between 16th and 19th century during the <a href="/wiki/Islamization" class="mw-redirect" title="Islamization">Islamization</a> of the Balkans.<a href="#cite_note-FOOTNOTEKovačević_et_al.2014-37">[36]</a> In the 2022 analysis Croatian, Bosnian and Herzegovinian, and Serbian individuals made "Northwestern Balkans" cluster which had less Southern European ("Greek") ancestry than "Northeastern Balkans" cluster formed by Romanian and Bulgarian individuals. The "NW Balkans" cluster also had "slightly longer within-cluster IBD segments than NE Balkans, which is matched with a consistently lower Ne and elevated <a href="/wiki/Runs_of_homozygosity" title="Runs of homozygosity">ROH</a>—suggestive of a smaller population than the northeast of the Balkans".<a href="#cite_note-FOOTNOTEGilbert_et_al.2022-60">[59]</a> According to a 2014 autosomal analysis of Western Balkan, the Croatian population shows genetic uniformity with other South Slavic populations. The Croatians and Bosnians were more close to East European populations and largely overlapped with Hungarians from Central Europe.<a href="#cite_note-FOOTNOTEKovačević_et_al.2014-37">[36]</a> In the 2015 analysis, they formed a western South Slavic cluster with the Bosnians and Slovenians in comparison to eastern cluster formed by Macedonians and Bulgarians with Serbians in the middle. The western cluster has an inclination toward Hungarians, Czechs, and Slovaks, while the eastern cluster toward Romanians and some extent Greeks.<a href="#cite_note-FOOTNOTEKushniarevich_et_al.2015-62">[61]</a> The modeled ancestral genetic component of Balto-Slavs among South Slavs was between 55 and 70%.<a href="#cite_note-FOOTNOTEKushniarevich_et_al.2015-62">[61]</a> In the 2018 analysis of <a href="/wiki/Slovenia" title="Slovenia">Slovenian</a> population, the Croatian population again clustered with Slovenians, Hungarians and was close to Czech. The population of Croatia mostly shares a common ancestry with Eastern, Western, and Southern Europeans, and has almost no relation to isolated populations like the <a href="/wiki/Sardinians" class="mw-redirect" title="Sardinians">Sardinians</a> and the <a href="/wiki/Basques" title="Basques">Basques</a>.<a href="#cite_note-FOOTNOTEKovačević_et_al.2014-37">[36]</a><a href="#cite_note-FOOTNOTEDelser_et_al.2018-63">[62]</a> According to 2023 archaeogenetic study autosomal qpAdm modelling, the modern-day Croats are 66.5% of Central-Eastern European early medieval Slavic ancestry, 31.2% of Croatia-Serbia local Roman ancestry and 2.4% Roman Imperial Era West Anatolian ancestry.<a href="#cite_note-Olalde2023-13">[13]</a> According to 2005 and 2006 autosomal STR marker studies the most isolated islands were Korčula,<a href="#cite_note-FOOTNOTEMartinović_Klarić_et_al.2005-64">[63]</a> and Susak.<a href="#cite_note-FOOTNOTEVitart_et_al.2006-42">[41]</a> A 2016 whole <a href="/wiki/Exome_sequencing" title="Exome sequencing">exome sequencing</a> study of 176 individuals from the <a href="/wiki/Vis_(island)" title="Vis (island)">island of Vis</a> confirmed the isolate status of the island's population, and revealed the "pattern of loss-of-function mutations, which resembles the trails of adaptive evolution".<a href="#cite_note-FOOTNOTEJeroncic_et_al.2016-65">[64]</a> The 2022 autosomal STR marker study on 2877 unrelated individuals from mainland (cities Zagreb, Pazin, Delnice, Zabok and Donji Miholjac, and region of Baranja) and insular (Krk, Cres, three North Dalmatian islands Ugljan-Pašman-Dugi Otok, Brač, Hvar, Korčula and Vis) subpopulations found higher genetic differentiation (0.005) compared to Southeastern Europe (0.002) indicating "a certain degree of genetic isolation, most likely due to the influence of endogamy within rural island populations". Croatian population is closest to Bosnia and Herzegovina, Hungary and Slovenia. There are visible four main clusters within sampled Croats, first including Mainland-Brač and Krk-North Dalmatian islands, second Hvar-Korčula, third Cres, and fourth Vis, the latter two having highest distances from the others.<a href="#cite_note-FOOTNOTENovokmet_et_al.2022a235–236,_241-66">[65]</a> <ul class="gallery mw-gallery-traditional center"> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px; height: 180px;"><a href="/wiki/File:Plos.Balkans.2.png" class="mw-file-description" title="Admixture analysis of autosomal SNPs in a global context on the resolution level of 7 assumed ancestral populations per Kovačević et al. (2014)"><img alt="Admixture analysis of autosomal SNPs in a global context on the resolution level of 7 assumed ancestral populations per Kovačević et al. (2014)" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/Plos.Balkans.2.png/150px-Plos.Balkans.2.png" decoding="async" width="150" height="59" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/Plos.Balkans.2.png/225px-Plos.Balkans.2.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a3/Plos.Balkans.2.png/300px-Plos.Balkans.2.png 2x" data-file-width="4033" data-file-height="1581" /></a></div> <div class="gallerytext">Admixture analysis of autosomal SNPs in a global context on the resolution level of 7 assumed ancestral populations per Kovačević et al. (2014)</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px; height: 180px;"><a href="/wiki/File:PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png" class="mw-file-description" title="Principal component (PC) analysis of the variation of autosomal SNPs in Western Balkan populations in Eurasian context per Kovačević et al. (2014)"><img alt="Principal component (PC) analysis of the variation of autosomal SNPs in Western Balkan populations in Eurasian context per Kovačević et al. (2014)" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6a/PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png/150px-PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png" decoding="async" width="150" height="99" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6a/PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png/225px-PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6a/PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png/300px-PCA_of_the_variation_of_autosomal_SNPs_in_Western_Balkan_populations_in_Eurasian_context.png 2x" data-file-width="3443" data-file-height="2283" /></a></div> <div class="gallerytext">Principal component (PC) analysis of the variation of autosomal SNPs in Western Balkan populations in Eurasian context per Kovačević et al. (2014)</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px; height: 180px;"><a href="/wiki/File:Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG" class="mw-file-description" title="Admixture analysis on the resolution level of 6 assumed ancestral populations per Kushniarevich et al. (2015)"><img alt="Admixture analysis on the resolution level of 6 assumed ancestral populations per Kushniarevich et al. (2015)" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG/150px-Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG" decoding="async" width="150" height="98" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG/225px-Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG/300px-Ancestral_components_of_the_Balto-Slavic_gene_pool.PNG 2x" data-file-width="2247" data-file-height="1461" /></a></div> <div class="gallerytext">Admixture analysis on the resolution level of 6 assumed ancestral populations per Kushniarevich et al. (2015)</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px; height: 180px;"><a href="/wiki/File:PLOS_3.PNG" class="mw-file-description" title="Genetic structure within European context according to three genetic systems atDNA (A), YDNA (B) and mtDNA (C) per Kushniarevich et al. (2015)"><img alt="Genetic structure within European context according to three genetic systems atDNA (A), YDNA (B) and mtDNA (C) per Kushniarevich et al. (2015)" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/PLOS_3.PNG/150px-PLOS_3.PNG" decoding="async" width="150" height="123" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/PLOS_3.PNG/225px-PLOS_3.PNG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/30/PLOS_3.PNG/300px-PLOS_3.PNG 2x" data-file-width="2250" data-file-height="1839" /></a></div> <div class="gallerytext">Genetic structure within European context according to three genetic systems atDNA (A), YDNA (B) and mtDNA (C) per Kushniarevich et al. (2015)</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px; height: 180px;"><a href="/wiki/File:PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png" class="mw-file-description" title="PC1vsPC2 plot based on whole genome SNP data per Kushniarevich et al. (2015)"><img alt="PC1vsPC2 plot based on whole genome SNP data per Kushniarevich et al. (2015)" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f5/PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png/126px-PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png" decoding="async" width="126" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f5/PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png/189px-PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f5/PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png/252px-PC1vsPC2_plot_based_on_whole_genome_SNP_data_of_Balto-Slavic_Speaking_Populations.png 2x" data-file-width="860" data-file-height="1022" /></a></div> <div class="gallerytext">PC1vsPC2 plot based on whole genome SNP data per Kushniarevich et al. (2015)</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=10" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 25em;"> <ul><li><a href="/wiki/White_Croats" title="White Croats">White Croats</a></li> <li><a href="/wiki/Origin_hypotheses_of_the_Croats" title="Origin hypotheses of the Croats">Origin hypotheses of the Croats</a></li> <li><a href="/wiki/Genetic_studies_on_Bosniaks" title="Genetic studies on Bosniaks">Genetic studies on Bosniaks</a></li> <li><a href="/wiki/Genetic_studies_on_Bulgarians" title="Genetic studies on Bulgarians">Genetic studies on Bulgarians</a></li> <li><a href="/wiki/Genetic_studies_on_Serbs" title="Genetic studies on Serbs">Genetic studies on Serbs</a></li> <li><a href="/wiki/Ancient_DNA" title="Ancient DNA">Ancient DNA</a></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Genetic history of Europe</a></li> <li><a href="/wiki/Y-DNA_haplogroups_by_ethnic_group" title="Y-DNA haplogroups by ethnic group">Y-DNA haplogroups by ethnic group</a></li> <li><a href="/wiki/Y-DNA_haplogroups_in_populations_of_Europe" title="Y-DNA haplogroups in populations of Europe">Y-DNA haplogroups in populations of Europe</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=11" title="Edit section: Notes">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol 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YFull.com. 10 November 2018. Retrieved 11 November 2018.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=I-P37+YTree+v6.07.08&rft.pub=YFull.com&rft.date=2018-11-10&rft_id=https%3A%2F%2Fwww.yfull.com%2Ftree%2FI-P37%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-27"><a href="#cite_ref-27">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.familytreedna.com/groups/i-2a-hap-group/about/results">"I2a Y-Haplogroup – Results: I2a2a-Dinaric"</a>. <a href="/wiki/Family_Tree_DNA" class="mw-redirect" title="Family Tree DNA">Family Tree DNA</a>. Retrieved 11 November 2018. <q>Ken Nordtvedt has split I2a2-M423-Dinaric into Din-N and Din-S. Din-N is older than Din-S. N=north of the Danube and S=south of the Danube River ... May 8, 2007: Dinaric I1b1 and DYS 448. DYS448 19 for S and 20 for N.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=I2a+Y-Haplogroup+%E2%80%93+Results%3A+I2a2a-Dinaric&rft.pub=Family+Tree+DNA&rft_id=https%3A%2F%2Fwww.familytreedna.com%2Fgroups%2Fi-2a-hap-group%2Fabout%2Fresults&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-28"><a href="#cite_ref-28">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBernie_Cullen2016" class="citation web cs1">Bernie Cullen (22 August 2016). <a rel="nofollow" class="external text" href="https://i2aproject.blogspot.com/2016/08/link-to-i-l621-tree-showing-major-str_22.html">"Link to I-L621 tree showing major STR clusters (Updated)"</a>. i2aproject.blogspot.com. <a href="/wiki/Blogger_(service)" title="Blogger (service)">Blogger</a>. Retrieved 3 April 2019.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=i2aproject.blogspot.com&rft.atitle=Link+to+I-L621+tree+showing+major+STR+clusters+%28Updated%29&rft.date=2016-08-22&rft.au=Bernie+Cullen&rft_id=https%3A%2F%2Fi2aproject.blogspot.com%2F2016%2F08%2Flink-to-i-l621-tree-showing-major-str_22.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-29"><a href="#cite_ref-29">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://docs.google.com/spreadsheets/d/1WkRe8UxrhMZ-CaYtPET5rIBbzqCNkGER4YJurbG2ESQ/edit#gid=198726360">"Y-DNA Haplogroup I and its Subclades – 2018"</a>. <a href="/wiki/ISOGG" class="mw-redirect" title="ISOGG">ISOGG</a>. 1 November 2018. Retrieved 11 November 2018.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Y-DNA+Haplogroup+I+and+its+Subclades+%E2%80%93+2018&rft.pub=ISOGG&rft.date=2018-11-01&rft_id=https%3A%2F%2Fdocs.google.com%2Fspreadsheets%2Fd%2F1WkRe8UxrhMZ-CaYtPET5rIBbzqCNkGER4YJurbG2ESQ%2Fedit%23gid%3D198726360&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-FOOTNOTEZupan_et_al.2013-31"><a href="#cite_ref-FOOTNOTEZupan_et_al.2013_31-0">^</a> <a href="#CITEREFZupan_et_al.2013">Zupan et al. 2013</a>. </li> <li id="cite_note-FOOTNOTEUtevska2017219–226,_302-32">^ <a href="#cite_ref-FOOTNOTEUtevska2017219–226,_302_32-0">a</a> <a href="#cite_ref-FOOTNOTEUtevska2017219–226,_302_32-1">b</a> <a href="#cite_ref-FOOTNOTEUtevska2017219–226,_302_32-2">c</a> <a href="#CITEREFUtevska2017">Utevska 2017</a>, p. 219–226, 302. </li> <li id="cite_note-FOOTNOTEŠarac_et_al.2016p._6:It_is_important_to_stress_that_the_proposed_old_age_of_the_I2a1b-M423_and_R1a1a1b1a*-M558_lineages_obtained_in_previous_studies_(Battaglia_et_al.,_2009;_Peričić_et_al.,_2005;_Rootsi_et_al.,_2004;_Underhill_et_al.,_2007,_2015)_has_been_based_on_STR_analysis_(8_and_10_loci,_respectively)_and_recent_studies_clearly_indicate_that_the_STR-based_age_calculations_tend_to_yield_overestimated_dates_(Batini_et_al.,_2015;_Hallast_et_al.,_2015;_Karmin_et_al.,_2015).-33"><a href="#cite_ref-FOOTNOTEŠarac_et_al.2016p._6:It_is_important_to_stress_that_the_proposed_old_age_of_the_I2a1b-M423_and_R1a1a1b1a*-M558_lineages_obtained_in_previous_studies_(Battaglia_et_al.,_2009;_Peričić_et_al.,_2005;_Rootsi_et_al.,_2004;_Underhill_et_al.,_2007,_2015)_has_been_based_on_STR_analysis_(8_and_10_loci,_respectively)_and_recent_studies_clearly_indicate_that_the_STR-based_age_calculations_tend_to_yield_overestimated_dates_(Batini_et_al.,_2015;_Hallast_et_al.,_2015;_Karmin_et_al.,_2015)._33-0">^</a> <a href="#CITEREFŠarac_et_al.2016">Šarac et al. 2016</a>, p. 6:It is important to stress that the proposed old age of the I2a1b-M423 and R1a1a1b1a*-M558 lineages obtained in previous studies (Battaglia et al., 2009; Peričić et al., 2005; Rootsi et al., 2004; Underhill et al., 2007, 2015) has been based on STR analysis (8 and 10 loci, respectively) and recent studies clearly indicate that the STR-based age calculations tend to yield overestimated dates (Batini et al., 2015; Hallast et al., 2015; Karmin et al., 2015).. </li> <li id="cite_note-34"><a href="#cite_ref-34">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBalanovsky2017" class="citation journal cs1">Balanovsky, O. (2017). <a rel="nofollow" class="external text" href="https://link.springer.com/journal/439/volumes-and-issues/136-5">"Toward a consensus on SNP and STR mutation rates on the human Y-chromosome (Review)"</a>. Human Genetics. 136 (5): 575–590. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00439-017-1805-8">10.1007/s00439-017-1805-8</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/28455625">28455625</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:253974249">253974249</a>. <q>While the reasons for the difference between genealogical and evolutionary Y-STR rates are thus partly understood, it remains unclear which rate to use. Many have applied the evolutionary rate, though quite a few have used the genealogical, or both, rates. Genetic genealogists generally apply the genealogical rate and criticize population-genetic studies for reporting (in their view) three-times overestimated ages ... The age of each haplogroup was also calculated using the STR genealogical rate and the STR evolutionary rate. Confidence intervals for the two STR-based ages (not shown on the plot) do not overlap. For example, the genealogical age of I2a-L621 (2200 ± 500 years) reaches the envelope age (from 2600 to 3100 ages), while the evolutionary age lies far beyond (9900 ± 2700 years). The observed pattern (Fig. 2a) clearly differs for haplogroups of different age classes. For ages less than 7000 years, the genealogical STR rate provides results consistent with or slightly underestimating the "true" ages, while the evolutionary rate results in three-fold overestimates. For ages between roughly 7000 and 15,000 years neither STR rate provides correct results. For haplogroups older than 15,000 years, the evolutionary rate estimates correctly or overestimates the "true" age.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Human+Genetics&rft.atitle=Toward+a+consensus+on+SNP+and+STR+mutation+rates+on+the+human+Y-chromosome+%28Review%29&rft.volume=136&rft.issue=5&rft.pages=575-590&rft.date=2017&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A253974249%23id-name%3DS2CID&rft_id=info%3Apmid%2F28455625&rft_id=info%3Adoi%2F10.1007%2Fs00439-017-1805-8&rft.aulast=Balanovsky&rft.aufirst=O.&rft_id=https%3A%2F%2Flink.springer.com%2Fjournal%2F439%2Fvolumes-and-issues%2F136-5&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-Fóthi-35">^ <a href="#cite_ref-Fóthi_35-0">a</a> <a href="#cite_ref-Fóthi_35-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFóthiGonzalezFehér2020" class="citation cs2">Fóthi, E.; Gonzalez, A.; Fehér, T.; et al. (2020), "Genetic analysis of male Hungarian Conquerors: European and Asian paternal lineages of the conquering Hungarian tribes", Archaeological and Anthropological Sciences, 12 (1), <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2020ArAnS..12...31F">2020ArAnS..12...31F</a>, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs12520-019-00996-0">10.1007/s12520-019-00996-0</a>, <q>We looked at 16 loci from 640 I2a-L621 samples in FTDNA's I2a project database and found that 7 individuals were 2 genetic steps away the Karos samples, of whom 1 was a Hungarian from Kunszentmárton, 2 were Ukrainians, 1 was Lithuanian, 1 was Belarusian, 1 was Russian, and 1 was a German from Poland. Based on SNP analysis, the CTS10228 group is 2200 ± 300 years old. The group's demographic expansion may have begun in Southeast Poland around that time, as carriers of the oldest subgroup are found there today. The group cannot solely be tied to the Slavs, because the proto-Slavic period was later, around 300–500 CE ... The SNP-based age of the Eastern European CTS10228 branch is 2200 ± 300 years old. The carriers of the most ancient subgroup live in Southeast Poland, and it is likely that the rapid demographic expansion which brought the marker to other regions in Europe began there. The largest demographic explosion occurred in the Balkans, where the subgroup is dominant in 50.5% of Croatians, 30.1% of Serbs, 31.4% of Montenegrins, and in about 20% of Albanians and Greeks. As a result, this subgroup is often called Dinaric. It is interesting that while it is dominant among modern Balkan peoples, this subgroup has not been present yet during the Roman period, as it is almost absent in Italy as well (see Online Resource 5; ESM_5). ... Their genetic haplogroup, I2a-CTS10228, is widespread among Slavs, but it is only present in 7% of Caucasian peoples, namely among the Karachay ... As such, it appears that the I2a-CTS10228 haplogroup in the paternal lineage of the Karos leaders arises from a specific branch in the Northern Caucasus dating to about 400–500 CE. Its modern descendents live among the Karachay, Hungarians, and various other surrounding nationalities.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Archaeological+and+Anthropological+Sciences&rft.atitle=Genetic+analysis+of+male+Hungarian+Conquerors%3A+European+and+Asian+paternal+lineages+of+the+conquering+Hungarian+tribes&rft.volume=12&rft.issue=1&rft.date=2020&rft_id=info%3Adoi%2F10.1007%2Fs12520-019-00996-0&rft_id=info%3Abibcode%2F2020ArAnS..12...31F&rft.aulast=F%C3%B3thi&rft.aufirst=E.&rft.au=Gonzalez%2C+A.&rft.au=Feh%C3%A9r%2C+T.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-HorolmaTibor2019-36"><a href="#cite_ref-HorolmaTibor2019_36-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPamjavFehérNémethKoppány_Csáji2019" class="citation book cs1 cs1-prop-foreign-lang-source">Pamjav, Horolma; Fehér, Tibor; Németh, Endre; Koppány Csáji, László (2019). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=xq2xDwAAQBAJ">Genetika és őstörténet</a> (in Hungarian). Napkút Kiadó. p. 58. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-963-263-855-3" title="Special:BookSources/978-963-263-855-3"><bdi>978-963-263-855-3</bdi></a>. <q>Az I2-CTS10228 (köznevén "dinári-kárpáti") alcsoport legkorábbi közös őse 2200 évvel ezelőttre tehető, így esetében nem arról van szó, hogy a mezolit népesség Kelet-Európában ilyen mértékben fennmaradt volna, hanem arról, hogy egy, a mezolit csoportoktól származó szűk család az európai vaskorban sikeresen integrálódott egy olyan társadalomba, amely hamarosan erőteljes demográfiai expanzióba kezdett. Ez is mutatja, hogy nem feltétlenül népek, mintsem családok sikerével, nemzetségek elterjedésével is számolnunk kell, és ezt a jelenlegi etnikai identitással összefüggésbe hozni lehetetlen. A csoport elterjedése alapján valószínűsíthető, hogy a szláv népek migrációjában vett részt, így válva az R1a-t követően a második legdominánsabb csoporttá a mai Kelet-Európában. Nyugat-Európából viszont teljes mértékben hiányzik, kivéve a kora középkorban szláv nyelvet beszélő keletnémet területeket.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Genetika+%C3%A9s+%C5%91st%C3%B6rt%C3%A9net&rft.pages=58&rft.pub=Napk%C3%BAt+Kiad%C3%B3&rft.date=2019&rft.isbn=978-963-263-855-3&rft.aulast=Pamjav&rft.aufirst=Horolma&rft.au=Feh%C3%A9r%2C+Tibor&rft.au=N%C3%A9meth%2C+Endre&rft.au=Kopp%C3%A1ny+Cs%C3%A1ji%2C+L%C3%A1szl%C3%B3&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dxq2xDwAAQBAJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"> </li> <li id="cite_note-FOOTNOTEKovačević_et_al.2014-37">^ <a href="#cite_ref-FOOTNOTEKovačević_et_al.2014_37-0">a</a> <a href="#cite_ref-FOOTNOTEKovačević_et_al.2014_37-1">b</a> <a href="#cite_ref-FOOTNOTEKovačević_et_al.2014_37-2">c</a> <a href="#cite_ref-FOOTNOTEKovačević_et_al.2014_37-3">d</a> <a href="#CITEREFKovačević_et_al.2014">Kovačević et al. 2014</a>. </li> <li id="cite_note-38"><a href="#cite_ref-38">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFUnderhill2015" class="citation cs2">Underhill, Peter A. 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id="cite_note-30"><a href="#cite_ref-30">^</a> The SNP I-P37 itself formed approximately 20 thousand <a href="/wiki/YBP" class="mw-redirect" title="YBP">YBP</a> and had <a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">TMRCA</a> 18 thousand YBP according to YFull,<a href="#cite_note-26">[26]</a> being too old and widespread as an SNP for argumentation of autochthony as well the old research used outdated nomenclature. According to "I-P37 (I2a)" project at <a href="/wiki/Family_Tree_DNA" class="mw-redirect" title="Family Tree DNA">Family Tree DNA</a>, the divergence at STR marker DYS448 20 > 19 is reported since 2007,<a href="#cite_note-27">[27]</a> while the SNP which defines the STR Dinaric-South cluster, I-PH908, is reported since 2014.<a href="#cite_note-28">[28]</a> The SNP I-PH908 at <a href="/wiki/ISOGG" class="mw-redirect" title="ISOGG">ISOGG</a> phylogenetic tree is named as I2a1a2b1a1a1c,<a href="#cite_note-29">[29]</a> while formed and had TMRCA approximately 1,800 YBP according to YFull.<a href="#cite_note-YFull-22">[22]</a> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=12" title="Edit section: References">edit</a>]</div> <dl><dt>Y-DNA</dt></dl> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBarać_et_al.2003" class="citation journal cs1">L. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Large-scale+migration+into+Britain+during+the+Middle+to+Late+Bronze+Age&rft.volume=601&rft.issue=7894&rft.pages=588-594&rft.date=2022&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8889665%23id-name%3DPMC&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A245509501%23id-name%3DS2CID&rft_id=info%3Abibcode%2F2022Natur.601..588P&rft_id=info%3Apmid%2F34937049&rft_id=info%3Adoi%2F10.1038%2Fs41586-021-04287-4&rft.aulast=Patterson&rft.aufirst=Nick&rft.au=Isakov%2C+Michael&rft.au=Booth%2C+Thomas&rft_id=https%3A%2F%2Fwww.nature.com%2Farticles%2Fs41586-021-04287-4_reference.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+studies+on+Croats" class="Z3988"></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Genetic_studies_on_Croats&action=edit&section=13" title="Edit section: External links">edit</a>]</div> <ul><li>The <a rel="nofollow" class="external text" href="https://www.familytreedna.com/groups/croatian-dna/about">Croatian DNA Project</a> at <a href="/wiki/Family_Tree_DNA" class="mw-redirect" title="Family Tree DNA">Family Tree DNA</a></li> <li>The <a rel="nofollow" class="external text" href="http://www.mefst.unist.hr/research/research-groups-and-laboratories/10-001-dalmatian-croatian-biobank/5038">10.001 Dalmatians – Croatian biobank</a> at <a href="/wiki/University_of_Split" title="University of Split">University of Split</a> School of Medicine</li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline 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Croatia">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/History_of_Croatia_before_the_Croats" title="History of Croatia before the Croats">Prehistoric</a></li> <li><a href="/wiki/Origin_hypotheses_of_the_Croats" title="Origin hypotheses of the Croats">Origins of Croats</a> <ul><li><a href="/wiki/White_Croats" title="White Croats">White Croats</a></li> <li><a href="/wiki/White_Croatia" title="White Croatia">White Croatia</a></li></ul></li> <li><a href="/wiki/Duchy_of_Croatia" title="Duchy of Croatia">Medieval duchy</a></li> <li><a href="/wiki/Dalmatian_city-states" title="Dalmatian city-states">Dalmatian city-states</a></li> <li><a href="/wiki/Slavs_in_Lower_Pannonia" title="Slavs in Lower Pannonia">Lower Pannonia</a></li> <li><a href="/wiki/Red_Croatia" title="Red Croatia">Red Croatia</a> <ul><li><a href="/wiki/Narentines" title="Narentines">Narentines</a></li> <li><a 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Provinces">Illyrian Provinces</a></li> <li><a href="/wiki/Kingdom_of_Croatia-Slavonia" title="Kingdom of Croatia-Slavonia">Austrio-Hungarian kingdom</a></li> <li><a href="/wiki/State_of_Slovenes,_Croats_and_Serbs" title="State of Slovenes, Croats and Serbs">World War I</a></li> <li><a href="/wiki/Kingdom_of_Yugoslavia" title="Kingdom of Yugoslavia">Kingdom of Yugoslavia</a> <ul><li><a href="/wiki/Banovina_of_Croatia" title="Banovina of Croatia">Banovina</a></li></ul></li> <li><a href="/wiki/World_War_II_in_Yugoslavia" title="World War II in Yugoslavia">World War II</a> <ul><li><a href="/wiki/Independent_State_of_Croatia" title="Independent State of Croatia">'Independent State'</a></li></ul></li> <li><a href="/wiki/Socialist_Republic_of_Croatia" title="Socialist Republic of Croatia">Socialist Republic</a></li> <li><a href="/wiki/Independence_of_Croatia" title="Independence of Croatia">Independence</a></li> <li><a href="/wiki/Croatian_War_of_Independence" title="Croatian War of Independence">War of Independence</a></li> <li><a href="/wiki/History_of_Croatia_(1995%E2%80%93present)" title="History of Croatia (1995–present)">Croatia since 1995</a></li> <li><a href="/wiki/Croatia%E2%80%93NATO_relations" title="Croatia–NATO relations">NATO</a></li> <li><a href="/wiki/2013_enlargement_of_the_European_Union" title="2013 enlargement of the European Union">European Union</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Geography_of_Croatia" title="Geography of Croatia">Geography</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Geography_of_Croatia#Climate" title="Geography of Croatia">Climate</a></li> <li><a href="/wiki/List_of_extreme_points_of_Croatia" title="List of extreme points of Croatia">Extreme points</a></li> <li><a href="/wiki/List_of_islands_of_Croatia" title="List of islands of Croatia">Islands</a></li> <li><a href="/wiki/List_of_lakes_of_Croatia" title="List of lakes of Croatia">Lakes</a></li> <li><a href="/wiki/List_of_mammals_of_Croatia" title="List of mammals of Croatia">Mammals</a></li> <li><a href="/wiki/List_of_mountains_in_Croatia" title="List of mountains in Croatia">Mountains</a></li> <li><a href="/wiki/List_of_protected_areas_of_Croatia" title="List of protected areas of Croatia">Protected areas</a></li> <li><a href="/wiki/List_of_rivers_of_Croatia" title="List of rivers of Croatia">Rivers</a></li> <li><a href="/wiki/Topography_of_Croatia" title="Topography of Croatia">Topography</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Politics_of_Croatia" title="Politics of Croatia">Politics</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Administrative_divisions_of_Croatia" title="Administrative divisions of Croatia">Administrative divisions</a></li> <li><a href="/wiki/Constitution_of_Croatia" title="Constitution of Croatia">Constitution</a></li> <li><a href="/wiki/President_of_Croatia" title="President of Croatia">President</a></li> <li><a href="/wiki/Government_of_Croatia" title="Government of Croatia">Government</a> <ul><li><a href="/wiki/Prime_Minister_of_Croatia" title="Prime Minister of Croatia">Prime Minister</a></li></ul></li> <li><a href="/wiki/Human_rights_in_Croatia" title="Human rights in Croatia">Human rights</a> <ul><li><a href="/wiki/LGBT_rights_in_Croatia" class="mw-redirect" title="LGBT rights in Croatia">LGBT</a></li></ul></li> <li><a href="/wiki/Law_enforcement_in_Croatia" title="Law enforcement in Croatia">Law enforcement</a></li> <li><a href="/wiki/Armed_Forces_of_Croatia" title="Armed Forces of Croatia">Military</a> <ul><li><a href="/wiki/General_Staff_of_the_Armed_Forces_(Croatia)" title="General Staff of the Armed Forces (Croatia)">General Staff</a></li> <li><a href="/wiki/Chief_of_the_General_Staff_of_the_Armed_Forces_(Croatia)" title="Chief of the General Staff of the Armed Forces (Croatia)">Chief of the General Staff</a></li></ul></li> <li><a href="/wiki/Croatian_Parliament" title="Croatian Parliament">Parliament</a> <ul><li><a href="/wiki/Speaker_of_the_Croatian_Parliament" title="Speaker of the Croatian Parliament">Speaker</a></li></ul></li> <li><a href="/wiki/List_of_political_parties_in_Croatia" title="List of political parties in Croatia">Political parties</a></li> <li><a href="/wiki/Elections_in_Croatia" title="Elections in Croatia">Elections</a></li> <li><a href="/wiki/Foreign_relations_of_Croatia" title="Foreign relations of Croatia">Foreign relations</a></li> <li><a href="/wiki/Croatian_security_and_intelligence_system" title="Croatian security and intelligence system">Security and intelligence</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Economy_of_Croatia" title="Economy of Croatia">Economy</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Croatian_brands" title="Croatian brands">Brands</a></li> <li><a href="/wiki/Energy_in_Croatia" title="Energy in Croatia">Energy</a></li> <li><a href="/wiki/List_of_Croatian_counties_by_GDP" title="List of Croatian counties by GDP">Gross domestic product (GDP)</a></li> <li><a href="/wiki/Industry_of_Croatia" title="Industry of Croatia">Industry</a></li> <li><a href="/wiki/History_of_Croatian_currency" title="History of Croatian currency">History of currency</a></li> <li><a href="/wiki/Croatian_National_Bank" title="Croatian National Bank">National Bank</a></li> <li><a href="/wiki/Croatia_and_the_euro" title="Croatia and the euro">The euro</a></li> <li><a href="/wiki/Privatization_in_Croatia" title="Privatization in Croatia">Privatization</a></li> <li><a href="/wiki/Zagreb_Stock_Exchange" title="Zagreb Stock Exchange">Stock Exchange</a></li> <li><a href="/wiki/Telecommunications_in_Croatia" title="Telecommunications in Croatia">Telecommunications</a></li> <li><a href="/wiki/Tourism_in_Croatia" title="Tourism in Croatia">Tourism</a></li> <li><a href="/wiki/Transport_in_Croatia" title="Transport in Croatia">Transport</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Category:Society_of_Croatia" title="Category:Society of Croatia">Society</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Croats" title="Croats">Croats</a></li> <li><a href="/wiki/Demographics_of_Croatia" title="Demographics of Croatia">Demographics</a></li> <li><a href="/wiki/Education_in_Croatia" title="Education in Croatia">Education</a></li> <li><a href="/wiki/Category:Ethnic_groups_in_Croatia" title="Category:Ethnic groups in Croatia">Ethnic groups</a></li> <li><a class="mw-selflink selflink">Genetics</a></li> <li><a href="/wiki/Healthcare_in_Croatia" title="Healthcare in Croatia">Healthcare</a></li> <li><a href="/wiki/Demographics_of_Croatia#Languages" title="Demographics of Croatia">Languages</a></li> <li><a href="/wiki/Religion_in_Croatia" title="Religion in Croatia">Religion</a></li> <li><a href="/wiki/Women_in_Croatia" title="Women in Croatia">Women</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;padding-left:0.5em;padding-right:0.5em;font-weight:normal;"><a href="/wiki/Culture_of_Croatia" title="Culture of Croatia">Culture</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Architecture_of_Croatia" title="Architecture of Croatia">Architecture</a></li> <li><a href="/wiki/Croatian_art" title="Croatian art">Art</a></li> <li><a href="/wiki/Cinema_of_Croatia" title="Cinema of Croatia">Cinema</a></li> <li><a href="/wiki/Croatian_cuisine" title="Croatian cuisine">Cuisine</a> <ul><li><a href="/wiki/Croatian_wine" title="Croatian wine">Wine</a></li></ul></li> <li><a href="/wiki/Croatian_language" title="Croatian language">Croatian language</a></li> <li><a href="/wiki/Croatian_literature" title="Croatian literature">Literature</a></li> <li><a href="/wiki/Music_of_Croatia" title="Music of Croatia">Music</a></li> <li><a href="/wiki/Public_holidays_in_Croatia" title="Public holidays in Croatia">Public holidays</a></li> <li><a href="/wiki/List_of_radio_stations_in_Croatia" title="List of radio stations in Croatia">Radio stations</a></li> <li><a href="/wiki/Sport_in_Croatia" title="Sport in Croatia">Sport</a></li> <li><a href="/wiki/Television_in_Croatia" title="Television in Croatia">Television</a></li> <li><a href="/wiki/List_of_World_Heritage_Sites_in_Croatia" title="List of World Heritage Sites in Croatia">World Heritage Sites</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;padding-left:0.5em;padding-right:0.5em;font-weight:normal;"><a href="/wiki/National_symbols_of_Croatia" title="National symbols of Croatia">Symbols</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lijepa_na%C5%A1a_domovino" title="Lijepa naša domovino">Anthem</a></li> <li><a href="/wiki/Croatian_checkerboard" title="Croatian checkerboard">Checkerboard</a></li> <li><a href="/wiki/Coat_of_arms_of_Croatia" title="Coat of arms of Croatia">Coat of arms</a></li> <li><a href="/wiki/Croatian_national_costume" title="Croatian national costume">Costume</a></li> <li><a href="/wiki/Crown_of_Zvonimir" title="Crown of Zvonimir">Crown</a></li> <li><a href="/wiki/Orders,_decorations,_and_medals_of_Croatia" title="Orders, decorations, and medals of Croatia">Decorations</a></li> <li><a href="/wiki/List_of_Croatian_flags" title="List of Croatian flags">Flags</a></li> <li><a href="/wiki/Croatian_interlace" title="Croatian interlace">Interlace</a></li> <li><a href="/wiki/List_of_Croatian_mottos" title="List of Croatian mottos">Motto</a></li> <li><a href="/wiki/Names_of_the_Croats_and_Croatia" title="Names of the Croats and Croatia">Names</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="font-weight:bold;"><div><div style="margin-bottom:-0.4em;"><ul><li><a href="/wiki/Outline_of_Croatia" title="Outline of Croatia">Outline</a></li><li><a href="/wiki/Index_of_Croatia-related_articles" title="Index of Croatia-related articles">Index</a></li></ul></div> <ul><li><a href="/wiki/Category:Croatia" title="Category:Croatia">Category</a></li> <li><a href="/wiki/Portal:Croatia" title="Portal:Croatia">Portal</a></li></ul></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Human_genetics" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Human_genetics" title="Template:Human genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Human_genetics" title="Template talk:Human genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Human_genetics" title="Special:EditPage/Template:Human genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Human_genetics" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_genetics" title="Human genetics">Human genetics</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Sub-topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_genome" title="Human genome">Human genome</a> <ul><li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li></ul></li> <li><a href="/wiki/Human_evolutionary_genetics" title="Human evolutionary genetics">Evolutionary genetics</a> <ul><li><a href="/wiki/Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">Human-chimp MRCA</a></li> <li><a href="/wiki/Neanderthal_genetics" title="Neanderthal genetics">Neanderthal genetics</a> <ul><li><a href="/wiki/Neanderthal_genome_project" title="Neanderthal genome project">Neanderthal genome project</a></li></ul></li> <li><a href="/wiki/Timeline_of_human_evolution" title="Timeline of human evolution">Timeline</a></li></ul></li> <li><a href="/wiki/Human_genetic_variation" title="Human genetic variation">Genetic variation</a> <ul><li><a href="/wiki/Blood_type_distribution_by_country" title="Blood type distribution by country">Blood type distribution by country</a></li> <li><a href="/wiki/Genealogical_DNA_test" title="Genealogical DNA test">Genealogical DNA test</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Race_and_genetics" title="Race and genetics">Race and genetics</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent evolution</a></li> <li><a href="/wiki/Surname_DNA_project" title="Surname DNA project">Surname DNA project</a></li></ul></li> <li><a href="/wiki/Human_genetic_enhancement" title="Human genetic enhancement">Genetic enhancement</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Genetic history</a> by region</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a> <ul><li><a href="/wiki/Genetic_history_of_North_Africa" title="Genetic history of North Africa">North Africa</a></li> <li><a href="/wiki/Genetic_history_of_West_Africa" title="Genetic history of West Africa">West Africa‎‎</a></li> <li><a href="/wiki/Genetic_history_of_Central_Africa" title="Genetic history of Central Africa">Central Africa</a></li> <li><a href="/wiki/Genetic_history_of_Eastern_Africa" title="Genetic history of Eastern Africa">Eastern Africa‎‎</a></li> <li><a href="/wiki/Genetic_history_of_Southern_Africa" title="Genetic history of Southern Africa">Southern Africa</a></li> <li><a href="/wiki/Genetic_history_of_the_African_diaspora" title="Genetic history of the African diaspora">African diaspora</a></li></ul></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a> <ul><li><a href="/wiki/Peopling_of_India" title="Peopling of India">India</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">Middle East</a> <ul><li><a href="/wiki/Early_Anatolian_farmers" class="mw-redirect" title="Early Anatolian farmers">Early Anatolian farmers</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_the_Caucasus" class="mw-redirect" title="Genetic history of the Caucasus">Caucasus</a> <ul><li><a href="/wiki/Caucasus_hunter-gatherer" title="Caucasus hunter-gatherer">Caucasus hunter-gatherer</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a> <ul><li><a href="/wiki/Early_European_Farmers" title="Early European Farmers">Early European Farmers</a></li> <li><a href="/wiki/Western_hunter-gatherer" title="Western hunter-gatherer">Western hunter-gatherer</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">British Isles</a></li> <li><a href="/wiki/Genetic_history_of_the_Iberian_Peninsula" title="Genetic history of the Iberian Peninsula">Iberia</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Eastern_hunter-gatherer" title="Eastern hunter-gatherer">Eastern hunter-gatherer</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Central_Asia" class="mw-redirect" title="Genetic history of Central Asia">Central Asia</a> <ul><li><a href="/wiki/Ancient_North_Eurasian" title="Ancient North Eurasian">Ancient North Eurasian</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_East_Asians" title="Genetic history of East Asians">East Asia</a> <ul><li><a href="/wiki/Ancient_Northeast_Asian" title="Ancient Northeast Asian">Ancient Northeast Asian</a></li> <li><a href="/wiki/Ancient_Paleo-Siberian" title="Ancient Paleo-Siberian">Ancient Paleo-Siberian</a></li> <li><a href="/wiki/Peopling_of_China" title="Peopling of China">China</a></li></ul></li> <li><a href="/wiki/Peopling_of_Southeast_Asia" title="Peopling of Southeast Asia">Southeast Asia</a> <ul><li><a href="/wiki/Peopling_of_Thailand" title="Peopling of Thailand">Thailand</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of Indigenous peoples of the Americas">America</a> <ul><li><a href="/wiki/Ancient_Beringian" title="Ancient Beringian">Ancient Beringian</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a> by group</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Europe <ul><li><a href="/wiki/Origin_of_the_Basques" title="Origin of the Basques">Basques</a></li> <li><a href="/wiki/Genetic_studies_on_Bosniaks" title="Genetic studies on Bosniaks">Bosniaks</a></li> <li><a href="/wiki/Genetic_studies_on_Bulgarians" title="Genetic studies on Bulgarians">Bulgarians</a></li> <li><a class="mw-selflink selflink">Croats</a></li> <li><a href="/wiki/Genetic_studies_on_Romanians" class="mw-redirect" title="Genetic studies on Romanians">Romanians</a></li> <li><a href="/wiki/Genetic_studies_on_Russians" title="Genetic studies on Russians">Russians</a></li> <li><a href="/wiki/Genetic_studies_on_Sami" title="Genetic studies on Sami">Sami</a></li> <li><a href="/wiki/Genetic_studies_on_Serbs" title="Genetic studies on Serbs">Serbs</a></li></ul></li> <li>MENA <ul><li><a href="/wiki/Genetic_studies_on_Arabs" title="Genetic studies on Arabs">Arabs</a></li> <li><a href="/wiki/Origin_of_the_Azerbaijanis" title="Origin of the Azerbaijanis">Azerbaijanis</a></li> <li><a href="/wiki/Genetic_history_of_Egypt" title="Genetic history of Egypt">Egyptians</a></li> <li><a href="/wiki/Genetic_studies_on_Jews" class="mw-redirect" title="Genetic studies on Jews">Jews</a></li> <li><a href="/wiki/Genetic_studies_on_Moroccans" title="Genetic studies on Moroccans">Moroccans</a></li> <li><a href="/wiki/Genetic_studies_on_Turkish_people" title="Genetic studies on Turkish people">Turks</a></li></ul></li> <li>South Asia <ul><li><a href="/wiki/Genetic_studies_on_Gujarati_people" title="Genetic studies on Gujarati people">Gujaratis</a></li> <li><a href="/wiki/Genetic_studies_on_Sinhalese" title="Genetic studies on Sinhalese">Sinhalese</a></li> <li><a href="/wiki/Genetic_studies_on_Sri_Lankan_Tamils" title="Genetic studies on Sri Lankan Tamils">Tamils (Sri Lankan)</a></li></ul></li> <li>East Asia <ul><li><a href="/wiki/Genetic_studies_on_Han_Chinese" class="mw-redirect" title="Genetic studies on Han Chinese">Han Chinese</a></li> <li><a href="/wiki/Genetic_and_anthropometric_studies_on_Japanese_people" title="Genetic and anthropometric studies on Japanese people">Japanese</a></li></ul></li> <li>Southeast Asia <ul><li><a href="/wiki/Genetic_studies_on_Filipinos" title="Genetic studies on Filipinos">Filipinos</a></li></ul></li> <li>Sub-Saharan Africa <ul><li><a href="/wiki/Genetic_studies_on_Hutu_and_Tutsi" class="mw-redirect" title="Genetic studies on Hutu and Tutsi">Hutu/Tutsi</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" 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