Croatian Croat Genetics: Abstracts and Summaries

by Kevin Alan Brook

Family Tree DNA - Genetic testing service
Family Tree DNA: Genetic Testing Service
DNA testing will show your connections with other families and ethnic groups. The database includes members of many Slavic and non-Slavic peoples from the Balkans. Croats are eligible to submit their paternal and maternal DNA data to Family Tree DNA's "Balkan Genetics" regional project as well as their other relevant regional project called the "Dinaric Alps DNA Project".

Croatia, located in the western Balkan Peninsula, shares borders with Slovenia (to its northwest), Hungary, Serbia (to its east), Bosnia-Herzegovina, and Montenegro and with the Adriatic Sea separating it from Italy. The Croatian language, similar to Serbian, belongs to the South Slavic language family.

Croats (Hrvati) are members of a specific ethnic group, while Croatians refers more to the nationality of all citizens of Croatia, although they are close enough to synonymous since Croats are around 90% of the country's population.

Croats descend from multiple ancient populations, including indigenous peoples like Illyrian tribes as well as immigrant Slavs and Middle Easterners.

Among members of Family Tree DNA's "Dinaric Alps DNA Project" project, their Y-DNA genealogical lines originating from Croatia include E-Z16988, I-M253, I-CTS10228, I-FGC22045, I-P37, I-S17250, I-Y3548, I-S17250, J-M205, N-M2019, R-M512, R-BY9003, R-Y2608, R-P278, and R-CTS3402.

Major studies of Croats

Jelena Šarac, Tena Šarić, Dubravka Havaš Auguštin, Natalija Novokmet, Nenad Vekarić, Mate Mustać, Blaženka Grahovac, Milijenko Kapović, Branimir Nevajda, Anton Glasnović, Saša Missoni, Siiri Rootsi, and Pavao Rudan. "Genetic heritage of Croatians in the Southeastern European gene pool-Y chromosome analysis of the Croatian continental and Island population." American Journal of Human Biology 28:6 (November 2016): pages 837-845. First published electronically on June 9, 2016.
      720 Croatian males had their Y-DNA examined here and compared with thousands of samples from different European ethnicities. Many different haplogroups were found among the Croats, yet 3 haplogroups stood out as particularly distinctive: I2a1b-M423 (found in over 30% of the Croatian samples and the most indigenous to the region), R1a1a1b1a*-M558 (found in 19% of the Croats and representing migrations of Slavs from central Europe), and E1b1b1a1b1a-V13 (found in about 9% of the Croats and stemming from farmers who arrived in Europe from the southeast).

Lovorka Barać, Marijana Peričić, Irena Martinović Klarić, Siiri Rootsi, Branka Janićijević, Toomas Kivisild, Jüri Parik, Igor Rudan, Richard Villems, and Pavao Rudan. "Y chromosomal heritage of Croatian population and its island isolates." European Journal of Human Genetics 11:7 (July 2013): pages 535-542.
      This study of the Y-DNA of 457 Croatians found haplogroup I to be particularly prevalent (38% frequency). R1a was found in 34% of these Croatians, and R1b found in 15%. Most Croatian lines descend from Paleolithic-era Europeans. Excerpts from the Abstract:

"Y chromosome variation in 457 Croatian samples was studied using 16 SNPs/indel and eight STR loci. High frequency of haplogroup I in Croatian populations and the phylogeographic pattern in its background STR diversity over Europe make Adriatic coast one likely source of the recolonization of Europe following the Last Glacial Maximum. The higher frequency of I in the southern island populations is contrasted with higher frequency of group R1a chromosomes in the northern island of Krk and in the mainland. R1a frequency, while low in Greeks and Albanians, is highest in Polish, Ukrainian and Russian populations and could be a sign of the Slavic impact in the Balkan region. Haplogroups J, G and E that can be related to the spread of farming characterize the minor part (12.5%) of the Croatian paternal lineages. In one of the southern island (Hvar) populations, we found a relatively high frequency (14%) of lineages belonging to P*(xM173) cluster, which is unusual for European populations. Interestingly, the same population also harbored mitochondrial haplogroup F that is virtually absent in European populations — indicating a connection with Central Asian populations, possibly the Avars."

Vincenza Battaglia, Simona Fornarino, Nadia Al-Zahery, Anna Olivieri, Maria Pala, Natalie M. Myres, Roy J. King, Siiri Rootsi, Damir Marjanović, Dragan Primorac, Rifat Hadžiselimović, Stojko Vidović, Katia Drobnič, Naser Durmishi, Antonio Torroni, Augusta Silvana Santachiara-Benerecetti, Peter A. Underhill, and Ornella Semino. "Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe." European Journal of Human Genetics 17:6 (June 2009): pages 820-830. First published online on December 24, 2008. (mirror)
      Y-DNA was collected and studied from 1206 males representing 17 populations, mostly from southeastern Europe. The Y-DNA haplogroup I was found in 45 percent of the Croats from Croatia included in this study. R1a was present in 27 percent of the Croat sample, and R1b was found among 13 percent of the Croats. Less than 15 percent of Croat men from Croatia belong to other Y-DNA haplogroups, and out of those most belong to haplogroup E which was found at a frequency of 9 percent among the Croat sample. 4.4 percent of the Croat sample belonged to J, 2 percent to N, and 1 percent to G.

Siiri Rootsi, Chiara Magri, Toomas Kivisild, Giorgia Benuzzi, Hela Help, Marina Bermisheva, Ildus Kutuev, Lovorka Barać, Marijana Peričić, Oleg Balanovsky, Andrey Pshenichnov, Daniel Dion, Monica Grobei, Lev A. Zhivotovsky, Vincenza Battaglia, Alessandro Achilli, Nadia Al-Zahery, Jüri Parik, Roy King, Cengiz Cinnioğlu, Elsa Khusnutdinova, Pavao Rudan, Elena Balanovska, Wolfgang Scheffrahn, Maya Simonescu, Antonio Brehm, Rita Goncalves, Alexandra Rosa, Jean-Paul Moisan, Andre Chaventre, Vladimir Ferak, Sandor Füredi, Peter J. Oefner, Peidong Shen, Lars Beckman, Ilia Mikerezi, Rifet Terzić, Dragan Primorac, Anne Cambon-Thomsen, Astrida Krumina, Antonio Torroni, Peter A. Underhill, A. Silvana Santachiara-Benerecetti, Richard Villems, and Ornella Semino. "Phylogeography of Y-Chromosome Haplogroup I Reveals Distinct Domains of Prehistoric Gene Flow in Europe". American Journal of Human Genetics 75:1 (2004): pages 128-137. Excerpt from the body of the article:

"A different scenario has to be envisioned for subhaplogroup I1b*, which is the most frequent clade in eastern Europe and the Balkans. It reaches its highest incidences in Croatia (31%) and Bosnia (40%), encompassing almost 80%-90% of I"

Marijana Peričić, Lovorka Barać-Lauc, Irena Martinović Klarić, Siiri Rootsi, Branka Janićijević, Igor Rudan, Rifet Terzić, Ivanka Čolak, Ante Kvesić, Dan Popović, et al. "High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations." Molecular Biology and Evolution 22:10 (October 2005): pages 1964-1975.
      This Y chromosomal study compares Croats with Slovenians, Serbs, Macedonian Slavs, Macedonian Greeks, Albanians, Poles, etc. Table 1 ("Summarized Percent Frequencies of R1b, R1a, I1b* (xM26), E3b1 and J2e)" reports that among 108 studied mainland Croatians their frequencies are 15.7% belonging to R1b-M173, 34.3% to R1a-M17 (but with a confidence range of 26% to 43.6%), 32.4% to I1b* (xM26) (but with a confidence range between 24.3% and 41.7%), 5.6% to E3b1-M78, and 1% to J2e-M102.

Dragan Primorac, Damir Marjanović, Pavao Rudan, Richard Villems, and Peter A. Underhill. "Croatian genetic heritage: Y-chromosome story." Croatian Medical Journal 52:3 (June 2011): pages 225-234.
      A summary of previous studies of Y-DNA evidence from Croats. They explain that more than 3/4 of Croatian men's Y-DNA lines started with early settlers of Europe while the remainder are descended from later arrivals coming in from the Middle East via southeastern Europe.

Emir Šehović, Martin Zieger, L. Spahić, D. Marjanović, and S. Dogan. "A glance of genetic relations in the Balkan populations utilizing network analysis based on in silico assigned Y-DNA haplogroups." Anthropological Review 81:3 (2018): pages 252-268. Published electronically on October 31, 2018.
      Table 3 lists the following Y-DNA haplogroup proportions among the study's "Croatian" samples: 36.1% I2a, 6.4% E1b1b, 23.8% R1a, 13.9% R1b, 3.9% I1, 0.9% J1, 1.9% J2a, 2.1% J2b.

Marijana Pericic Salihovic, Lovorka Barać-Lauc, Irena Martinović Klarić, and Pavao Rudan. "Review of Croatian genetic heritage as revealed by mitochondrial DNA and Y chromosomal lineages." Croatian Medical Journal 46:4 (September 2005): pages 502-513.
      This summary of the previously collected results of 721 people's mtDNA and Y-DNA haplogroups includes both mainland Croatians and insular Croatians (living on islands and islets). This includes 451 men's Y-DNA. Excerpts from the Abstract:

[...] The phylogeography of mtDNA and Y chromosome variants of Croatians can be adequately explained within typical European maternal and paternal genetic landscape, with the exception of mtDNA haplogroup F and Y-chromosomal haplogroup P* which indicate a connection to Asian populations. Similar to other European and Near Eastern populations, the most frequent mtDNA haplogroups in Croatians were H (41.1%), U5 (10.3%), and J (9.7%). The most frequent Y chromosomal haplogroups in Croatians, I-P37 (41.7%) and R1a-SRY1532 (25%), as well as the observed structuring of Y chromosomal variance reveal a clearly evident Slavic component in the paternal gene pool of contemporary Croatian men. [...]

Pierpaolo Maisano Delser, Metka Ravnik-Glavač, Paolo Gasparini, Damjan Glavač, and Massimo Mezzavilla. "Genetic Landscape of Slovenians: Past Admixture and Natural Selection Pattern." Frontiers in Genetics (November 19, 2018).
      Autosomal clustering tools, such as principal component analysis (PCA), show that some Croatians genetically cluster with Slovenians, Hungarians, and Czechs in what the researchers call a "Central-Eastern European" cluster, but other Croatians group with Sicilians, Greeks, Romanians, and Northern Italians in a "Southern European" cluster.

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