Haplogroup I-M438 Explained

I-M438
Origin-Date:	28–33,000 years ago
Origin-Place:	South-Eastern or Eastern Europe.
Ancestor:	I-M170
Descendants:	I-L460, I-L1251
Mutations:	M438/P215/S31
Members:	I2a1a: Sardinia^[1] I2a1b: Bosnia and Herzegovina,^[2] I2a2: Britain, Germany, and Sweden

Haplogroup I-M438, also known as I2 (ISOGG 2019), is a human DNA Y-chromosome haplogroup, a subclade of haplogroup I-M170. Haplogroup I-M438 originated some time around 26,000–31,000 BCE. It originated in Europe and developed into several main subgroups: I2-M438*, I2a-L460, I2b-L415 and I2c-L596.^[1] The haplogroup can be found all over Europe and reaches its maximum frequency in the Dinaric Alps (Balkans) via founder effect, related to the migrations of the Early Slavs to the Balkan peninsula.

The oldest example so far found is that of Hohle Fels (49) from Germany, being at least 14,200 years old.^[3]

Origin and prehistoric presence

Haplogroup I2a was the most frequent Y-DNA among Mesolithic Western European hunter-gatherers (WHG) belonging to Villabruna Cluster. A 2015 study found haplogroup I2a in 13,500 year old remains from the Azilian culture (from Grotte du Bichon, modern Switzerland).^[4] Subclades of I2a1 (I-P37.2), namely I-M423 and I-M26, have been found in remains of WHGs dating from 10,000 to 8,000 years before present.^[5]

In a 2015 study published in Nature, the remains of six individuals from Motala ascribed to the Kongemose culture were successfully analyzed. With regards to Y-DNA, two individuals were ascribed to haplogroup I2a1b, one individual was ascribed to haplogroup I2a1, and one individual was ascribed to haplogroup I2c.^[6]

Subclades of I-L460

I-P37.2

The I-P37.2+ (also known as I2a1a) (ISOGG 2019). The subclade divergence for I-P37.2 occurred 10.7±4.8 kya. The age of YSTR variation for the P37.2 subclade is 8.0±4.0 kya.^[1] It is the predominant version of I2 in Eastern Europe.^[7] The I2a is further made up by subgroups I-M26, I-M423, I-L1286, I-L880.

I-L158

Haplogroup I-M26 (or M26) I2a1a1a (ISOGG 2019).

Haplogroup I-L158 (L158, L159.1/S169.1, M26) accounts for approximately 40% of all patrilines among Sardinians.^[8] ^[9] It is also found at low to moderate frequency among populations of the Pyrenees (9.5% in Bortzerriak, Navarra; 9.7% in Chazetania, Aragon; 8% in Val d'Aran, Catalunya; 2.9% in Alt Urgell, Catalunya; and 8.1% in Baixa Cerdanya, Catalunya) and Iberia, and it has been found in 1.6% of a sample of Albanians living in the Republic of North Macedonia^[10] and 1.2% (3/257) of a sample of Czechs.^[11] The age of YSTR variation for the M26 subclade has been calculated as 8.0±4.0 kya.^[1]

I-L178

I-L178 is very rare, but has been found in two persons from Germany and one from Poland. The age of YSTR variation for the M423 subclade is 8.8±3.6 kya.^[12]

I2a-L621

I2a1a2b-L621 is typical of the Slavic populations, being highest in Southeastern European regions of Bosnia-Herzegovina and South Croatia (>45%),^[2] ^[13] ^[14] in Croats (37.7-69.8%), Bosniaks (43.53-52.17%), and Serbs (36.6-42%)often called "Dinaric".^[15] It has the highest variance and concentration in Eastern Europe (i.e., Ukraine, Southeastern Poland, Belarus).^[16] According to YFull YTree it formed 11,400 YBP and had TMRCA 6,500 YBP, while its main subclades lineage is 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-Y18331 (2,100 YBP) / I-Z17855 (2,100-1650 YBP) / I-Y4460 (2,100 YBP) / I-S17250 (2,100-1,850 YBP) > I-PH908 (1,850-1,700 YBP).^[17]

Older research considered that the high frequency of this subclade in the South Slavic-speaking populations to be the result of "pre-Slavic" paleolithic settlement in the region. Peričić et al. (2005) for instance placed its expansion to have occurred "not earlier than the YD to Holocene transition and not later than the early Neolithic".^[1] ^[2] ^[18] ^[19] However, the prehistoric autochthonous origin of the haplogroup I2 in the Balkans is now considered as outdated, as already Battaglia et al. (2009) observed highest variance of the haplogroup in Ukraine, and Zupan et al. (2013) noted that it suggests it arrived with Slavic migration from the homeland which was in present-day Ukraine.^[20] O.M. Utevska (2017), in her PhD thesis, despite being a part of research team who came to a different conclusion in 2015,^[21] proposed that the haplogroup STR haplotypes have the highest diversity in Ukraine, with ancestral STR marker result "DYS448=20" comprising "Dnieper-Carpathian" cluster, while younger derived result "DYS448=19" comprising the "Balkan cluster" which is predominant among the South Slavs.^[16] According to her, this "Balkan cluster" also has the highest variance in Ukraine, which indicates that the very high frequency in the Western Balkan is probably because of a founder effect.^[16] Utevska calculated that the STR cluster divergence and its secondary expansion from the middle reaches of the Dnieper river or from Eastern Carpathians 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 Cucuteni–Trypillia culture.^[16] However, STR-based calculations give overestimated dates,^[22] ^[23] and more specifically, the "Balkan cluster" is represented by a single SNP, I-PH908, known as I2a1a2b1a1a1c in ISOGG phylogenetic tree (2019), and according to YFull YTree it formed and had TMRCA approximately 1,850-1,700 YBP (2nd-3rd century AD).^[17]

It is considered that I-L621 could have been present in the Cucuteni–Trypillia culture,^[24] but until now was mainly found G2a and non-I2-L621 clades,^[25] ^[26] and another clade I2a1a1-CTS595 was present in the Baden culture of the Chalcolithic Carpathian Basin.^[24] ^[27] ^[28] Although it is dominant among the modern Slavic peoples on the territory of the former Balkan provinces of the Roman Empire, until now it was not found among the samples from the Roman period and is almost absent in contemporary population of Italy.^[15] 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 Poland, is mainly related to the Slavs and their medieval migration, and the "largest demographic explosion occurred in the Balkans".^[15] ^[29] According to a 2023 archaeogenetic study, I2a-L621 is absent in the antiquity and appears only since the Early Middle Ages "always associated with Eastern European related ancestry in the autosomal genome, which supports that these lineages were introduced in the Balkans by Eastern European migrants during the Early Medieval period."^[30]

Some of the earliest archeogenetic samples until now is Sungir 6 (~900 YBP) near Vladimir, Russia which belonged to the I-S17250 > I-Y5596 > I-Z16971 > I-Y5595 > I-A16681 subclade,^[31] ^[32] as well I-CTS10228 and I-Y3120 subclades found in two Vikings from Sweden (VK53) and Ukraine (VK542) with predominantly Slavic ancestry of which the second belongs to Gleb Svyatoslavich (11th century).^[33] It was also found in the skeletal remains of Hungarian conquerors of the Carpathian Basin from the 9th century, part of Western Eurasian-Slavic component of the Hungarians.^[15] ^[24] ^[34]

I-M223

Haplogroup I-M223 aka I2a1b1 (ISOGG 2019), formerly I2a2a (ISOGG 2014). The age of YSTR variation for the I-M223 subclade has been variously estimated as 13.2±2.7 kya,^[1] 12.3±3.1 kya.,^[12] 14.6 kya^[35] and 14.6±3.8 kya (Rootsi 2004). I-M223 has a peak in Germany and another in the northeast of Sweden, but also appears in Romania/Moldova, Russia, Greece, Italy and around the Black Sea.^[36] Haplogroup I-M223 has been found in over 4% of the population only in Germany, the Netherlands, Belgium, Denmark, Scotland, and England (excluding Cornwall) – also the southern tips of Sweden and Norway in Northwest Europe; the provinces of Normandy, Maine, Anjou, and Perche in northwestern France; the province of Provence in southeastern France; the regions of Tuscany, Umbria, and Latium in Italy; Moldavia and the area around Russia's Ryazan Oblast and Mordovia in Eastern Europe. Of historical note, both haplogroups I-M253 and I-M223 appear at a low frequency in the historical regions of Bithynia and Galatia in Turkey. Haplogroup I-M223 also occurs among approximately 1% of Sardinians.

I-M284

Haplogroup I2a1b1a1a (ISOGG 2019) or I-M284, has been found almost exclusively amongst the populations of the United Kingdom and Ireland suggesting that it may have arisen amongst the Ancient Britons, with a most recent common ancestor (MRCA) who lived about 3,100 years BP.^[37] The presence of this subclade "provides some tentative evidence of ancient flow with eastern areas that could support the idea that the [late Celtic] La Tene culture was accompanied by some migration."

Where it is found in those of predominately Irish descent, with Gaelic surnames, it may suggest an ancestor who arrived in Ireland during prehistory, from Celtic Britain.^[38] For example, I-M284 includes many males with the surnames McGuinness and McCartan, who have a single, historically recorded male ancestor in the 6th century; thus it is unlikely to be the result of subsequent migration from Britain to Ireland.^[38] Some subclades of I-M284 that are atypical of Ireland are relatively common in continental Europe,^[38] which also supports a point of origin east of Ireland.

I-CTS10057

Continentals. Mother Haplogroup for group I-Z161 (Continental 1 and 2) and I-L701 group (Continental 3). Around 10.000 years old.

I-Z161

Haplogroup I2a1b1a2b (ISOGG 2019). Z161+ defines the I2 Continental clade 1 and 2. Its age is estimated around 7,000 years old. It is mainly found in North Europe, especially in Denmark, Germany, the Netherlands, and England. In Northwest Sicily it can also be found; this is believed to be due to remnants of a Norman settlement.

I-L701

Called Continental 3. Continental 3 has a wide distribution. Found in Central Europe from Germany, Austria to Poland, Romania and Ukraine, but also in lower frequencies in Greece, Italy, France, Spain, England, Ireland, and Armenia. It may have been disseminated in part by the Goths. It is nearly absent from Scandinavia and Scotland.

I-M436

Subclades list

Up-to-date phylogenetic trees listing subclades of I can be found at Y-Full and FamilyTreeDNA

•I2 M438/P215/PF3853/S31 (16638804 A->G)
• •I2a L460/PF3647/S238 (7879415 A->C)
• • •I2a1 P37.2/PF4004 (14491684 T->C)
• • • •I2a1a CTS595 (6874115 C->T)
• • • • •I2a1a1 M26/PF4056 (21865821 G->A)
• • • • • •I2a1a1a CTS8968/PF4036 (18584762 T->C)
• • • • • • •I2a1a1a1 L672/S327 (22228628 T->A)
• • • • • • • •I2a1a1a1a L160/PF4013/S184
• • • • • • •I2a1a1a2 CTS6027 (16620529 C->G) w. Europeans
• • • • • • •I2a1a1a3 Z27395 (16681589 A->G)
• • • • • • • •I2a1a1a3a Z27410 (18891640 T->G)
• • • • • • • • •I2a1a1a3a1 Z27430 (15383072 T->C)
• • • • • • • • • •I2a1a1a3a1a Z27446 (14420039 G->A)
• • • • • • • • • • •I2a1a1a3a1a1 Z27447 (17445510 A->G)
• • • • • • • • • • • •I2a1a1a3a1a1a Z27452 (7346620 C->T) Sardinians
• • • • • • • • • • • •I2a1a1a3a1a1b Z27453 (22045753 C->A)
• • • • • • • • • • • • •I2a1a1a3a1a1b1 Z27456 (13885862 T->C) Sardinians
• • • • • •I2a1a1b Z27354 (7132292 T->C)
• • • • • • •I2a1a1b1 Z27400 (2833818 C->T) Sardinians
• • • • •I2a1a2 S21825 (19126655 G->A)
• • • • • •I2a1a2a L1286 (21778662 G->A)
• • • • • • •I2a1a2a1 L1287 (21970862 G->T)
• • • • • • • •I2a1a2a1a L233/S183 (14487362 G->A)
• • • • • • • • •I2a1a2a1a1 A417 (8846964 T->C) Brits
• • • • • • •I2a1a2a2 L880 (3436270 C->T) central Europeans
• • • • • •I2a1a2b L1294 (2887401 T->C) French
• • • • •I2a1a3 S2699 (14434372 C->T)
• • • •I2a1b M423 (19096091 G->A)
• • • • •I2a1b1 L161.1/S185.1 (22513718 C->T)
• • • • • •I2a1b1a L1498/Y4008 (18668472 C->T)
• • • • • • •I2a1b1a1 FGC7218/Y5280 (21354226 G->A) Irish
• • • • • • •I2a1b1a2 FGC14448/Y5450 (7158983 A->G) Irish
• • • • • • •I2a1b1a3 FGC7108/S2640 (14107847 G->A)
• • • • • • • •I2a1b1a3a FGC7197/S7703 (17494645 G->C)
• • • • • • • • •I2a1b1a3a1 S7714 (21345894 T->A)
• • • • • • • • • •I2a1b1a3a1a S8889 (4310132 A->G)
• • • • • • • •I2a1b1a3b S2742 (19440521 A->G)
• • • • • • • •I2a1b1a3c FGC7173/Y3729 (8472670 A->T)
• • • • • • • • •I2a1b1a3c1 FGC7156/Y4665 (2685333 T->C)
• • • • • • • • • •I2a1b1a3c1a FGC7218/Y5280 (21354226 G->A)
• • • • • • • • •I2a1b1a3c2 FGC14453/Y5456 (16637771 A->G)
• • • • • • •I2a1b1a4 A1150 (15543414 A->T)
• • • • • •I2a1b1b PF4135 (18981938 G->A) Brits
• • • • • •I2a1b1c CTS8849.2 (18236845 A->G)
• • • • • •I2a1b1d PF6316.2 (18181850 T->G)
• • • • •I2a1b2 L621/S392 (18760081 G->A)
• • • • • •I2a1b2a CTS4002 (15328797 C->A)
• • • • • • •I2a1b2a1 CTS10228 (9334357 A->C)
• • • • • • • •I2a1b2a1a S17250/YP204 (15531354 G->A)
• • • • • • • • •I2a1b2a1a1 Z16971 (6676628 G->C) Slovenians, Germans, Ukrainians
• • • • • • • • •I2a1b2a1a2 Y3548 (14856842 A->G) Sardinians
• • • • • • • • • •I2a1b2a1a2a Y4882 (21135222 G->A) Poles
• • • • • • • • •I2a1b2a1a3 A356/Z16983 (17558968 T->C)
• • • • • • • • • •I2a1b2a1a3a Y6651 (23792119 A->G) Czechs
• • • • • • • •I2a1b2a1b Y4460 (9028830 G->A)
• • • • • • • • •I2a1b2a1b1 Y3118 (23548545 T->C)
• • • • • • • • • •I2a1b2a1b1a CTS5579/S5054 (16378157 G->T) Finns
• • • • • • • •I2a1b2a1b V19.2 (2189,200 T->C)
• • • • • • • •I2a1b2a1c P61.2 (14482194 G->A)
• • •I2a2 M436/P214/PF3856/S33 (18747493 G->C)
• • • •I2a2a M223 (21717307 G->A)
• • • • •I2a2a1 CTS616 (6906332 G->C)
• • • • • •I2a2a1a Y3721 (7982615 C->T)
• • • • • • •I2a2a1a1 M284 (22750461..22750464 ACAA->del)
• • • • • • • •I2a2a1a1a L1195 (18865320 G->A)
• • • • • • • • •I2a2a1a1a1 L126/S165 (14901633 C->T)
• • • • • • • • • •I2a2a1a1a1a FGC20048/S7753/Y4171 (17603051 G->T)
• • • • • • • • • • •I2a2a1a1a1a1 FGC20065/Y4751 (18635722 A->G)
• • • • • • • • • • • •I2a2a1a1a1a1a Y4750 (23059262 C->G)
• • • • • • • • • • • • •I2a2a1a1a1a1a1 FGC20062/Y4753 (16910733 G->T) Irish, Scots
• • • • • • • • • • • • •I2a2a1a1a1a1a2 Y8599 Irish
• • • • • • • • • •I2a2a1a1a1b Y7190 (7987909 C->T) Scots
• • • • • • • • • •I2a2a1a1a1c F3878.2 (8752246 G->A)
• • • • • • • • •I2a2a1a1a2 L1193 (9448484 C->A)
• • • • • • • • • •I2a2a1a1a2a CTS4922 (15903939 T->C)
• • • • • • • • • • •I2a2a1a1a2a1 FGC14237/Y5996 (8292320 G->T) Irish
• • • • • • • • • •I2a2a1a1a2b CTS3792/S2380 (15183633 C->T)
• • • • • • •I2a2a1a2 Z2057 (4770006 T->C)
• • • • • • • •I2a2a1a2a L1229 (14937828 C->A)
• • • • • • • • •I2a2a1a2a1 Y3681 (22519059 T->C)
• • • • • • • • • •I2a2a1a2a1a Z2058 (5317533 T->C)
• • • • • • • • • • •I2a2a1a2a1a1 Z2054 (2980725 C->T)
• • • • • • • • • • • •I2a2a1a2a1a1a BY524/Y9443 (6371561 A->G)
• • • • • • • • • • • • •I2a2a1a2a1a1a1 L812/S391 (14850035 G->A)
• • • • • • • • • • • • • •I2a2a1a2a1a1a1a Y5308 (16820196 G->A)
• • • • • • • • • • • • •I2a2a1a2a1a1a2 Y10648 (8267867 C->G)
• • • • • • • • • • • •I2a2a1a2a1a1b Y7244 (8271590 C->T) Germans
• • • • • • • • • • • • •I2a2a1a2a1a1b1 P53.3 (14491649 T->C)
• • • • • • • • • • • • •I2a2a1a2a1a1b2 Y7243 (2911078 G->A)
• • • • • • • • • • • •I2a2a1a2a1a1c FGC15106/Y4334 (7845669 G->T)
• • • • • • • • • • • • •I2a2a1a2a1a1c1 FGC15128/Y4714 (15945966 G->C) Irish
• • • • • • • • • • • • •I2a2a1a2a1a1c2 Y4760 (15025945 A->T) Brits, Germans
• • • • • • • • • • • •I2a2a1a2a1a1d BY138 (2809424 A->T) Mexicans, English
• • • • • • • • •I2a2a1a2a2 S18331 (16449714 C->T) Brits
• • • • • • • • • •I2a2a1a2a2a L1230 (3436442 G->A) Brits
• • • • • • • •I2a2a1a2b Y7240 (2688254 T->G) English
• • • • • •I2a2a1b CTS10057 (19232160 C->T)
• • • • • • •I2a2a1b1 L702 (7629205 C->T)
• • • • • • • •I2a2a1b1a P78 (6740387 G->A)
• • • • • • • • •I2a2a1b1a1 S25733 (23595728 C->T)
• • • • • • • • • •I2a2a1b1a1a A427 (23281929 G->T)
• • • • • • • • • • •I2a2a1b1a1a1 S23612 (21826246 C->T) Brits, Italians, Germans
• • • • • • • • • • • •I2a2a1b1a1a1a Y5360 (23185624 G->T)
• • • • • • • • • • • •I2a2a1b1a1a1b S10702 (7822881 T->C)
• • • • • • • • • • •I2a2a1b1a1a2 Y5369 (23185624 G->T) Brits
• • • • • • • • •I2a2a1b1a2 Y7219 (8447173 C->A)
• • • • • • • • • •I2a2a1b1a2a Y7214 (4901002 C->A) Dutch
• • • • • • • • • •I2a2a1b1a2b Y8945 (4567154 G->A)
• • • • • • • •I2a2a1b1b L699 (2663920 A->G)
• • • • • • • • •I2a2a1b1b1 L704 (17595966 C->A)
• • • • • • • • • •I2a2a1b1b1a S12195 (9105096 C->T)
• • • • • • • • • • •I2a2a1b1b1a1 Y6973 (8430276 A->G)
• • • • • • • • •I2a2a1b1b2 Z26416 (18851267 G->A) Sardinians
• • • • • • • • •I2a2a1b1b3 PF6901 (17558585 C->G) Sardinians
• • • • • • • •I2a2a1b1c PF6896 (8904322 C->T) Brits, French
• • • • • • •I2a2a1b2 Z161 (2696497 C->G)
• • • • • • • •I2a2a1b2a L801/S390 (21763755 A->C)
• • • • • • • • • •I2a2a1b2a1 CTS1977 (14140273 G->A)
• • • • • • • • • • •I2a2a1b2a1a Y5282 (7982615 C->T)
• • • • • • • • • • • •I2a2a1b2a1a1 S8522 (3162610 T->C)
• • • • • • • • • • • • •I2a2a1b2a1a1a P95 (14869706 G->T) Poles, n.w. Europeans
• • • • • • • • • • • •I2a2a1b2a1a2 CTS1858 (14098206 G->T)
• • • • • • • • • • • • •I2a2a1b2a1a2a CTS10148 (19291083 A->T) African Americans
• • • • • • • • • • •I2a2a1b2a1b Y7152 (5852771 C->T)
• • • • • • • • • • •I2a2a1b2a1c BY526 or Y8935
• • • • • • • • • •I2a2a1b2a2 CTS6433 (16889964 T->C)
• • • • • • • • • • •I2a2a1b2a2a S2364 (23052606 A->G)
• • • • • • • • • • • •I2a2a1b2a2a1 FGC3618/S2361 (17355245 A->G)
• • • • • • • • • • • • •I2a2a1b2a2a1a Z78 (9154908 T->C)
• • • • • • • • • • • • • •I2a2a1b2a2a1a1 CTS8584
• • • • • • • • • • • • • • •I2a2a1b2a2a1a1a Z185 (22929384 T->C)
• • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1 Z180 (22065434 A->G)
• • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a L1198 (18405788 C->T))
• • • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a1 Z190 (17473966 G->T)
• • • • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a1a S434/Z79 (17147721 G->A) English
• • • • • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a1a1 Y5729 (14394460 C->T) English
• • • • • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a1a2 Y7682 (not yet reported)
• • • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a1b Y7280 (17305030 T->G)
• • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a2 FGC3617/S2368/Z166 (14010863 G->T)
• • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1a3 P195.2 (22665262 A->G) Brits
• • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1b Y5748 (14106278 C->T)
• • • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1b1 Y7272 (19264274 G->A)
• • • • • • • • • • • • • • • •I2a2a1b2a1b1a1a1c Y7273 (16449864 A>G)
• • • • • • • • • • • • •I2a2a1b2a1b1a2 Y9161 (9154908 T->C)
• • • • • • • • • • • •I2a2a1b2a1b1b Y4925/ZS20 (23977120 A->C)
• • • • • • • • • • • • •I2a2a1b2a1b1b1 CTS661/L1272 (6931960 G->A)
• • • • • • • • • • • • • •I2a2a1b2a1b1b1a Y5717 (6931960 G->A)
• • • • • • • • • • • • •I2a2a1b2a1b1b2 FGC17399/Y4926 (23549293 G->A)
• • • • • • • • • • • • • •I2a2a1b2a1b1b2a S8104 (7432604 A->G)
• • • • • • • • • • • •I2a2a1b2a1b1c Y4955 (15362162 G->A)
• • • • • • • • • • • • •I2a2a1b2a1b1c1 Y5695 (14574229 T->C)
• • • • • • • • • • • • • •I2a2a1b2a1b1c1a Y7263 (7900806 C->A)
• • • • • • • • • • • • • • •I2a2a1b2a1b1c1a1 Y7265 (9147689 T->C)
• • • • • • • • • • • • • •I2a2a1b2a1b1c1b FGC20004/Y5692 (18993826 A->T)
• • • • • • • • • • • • •I2a2a1b2a1b1c2 Y10659 (4616653 A->G)
• • • • • • • • • • •I2a2a1b2a1b2 L1425 (14491835 C->T)
• • • • • • • • • • • •I2a2a1b2a1b2a CTS5332.2/PF7472.2 (16212441 G->A)
• • • • • • • • • • • •I2a2a1b2a1b2b Y4769 (18201094 A->G) Germans
• • • • • • • • • • •I2a2a1b2a1b3 Y7426 (18830768 A->G) English
• • • • • • • • • • •I2a2a1b2a1b4 S243/Z63 (14401486 C->T)
• • • • • • • • • • •I2a2a1b2a1b5 M3268.2/PF3292.2 (8519704 G->A) Germans
• • • • • • • • • • •I2a2a1b2a1b6 DF5.2/S191.2 (18393814 A->G) Brits
• • • • • • • • • • •I2a2a1b2a1b7 F3406.2 (23119516 G->A)
• • • • • • • • • •I2a2a1b2a2 L1290 (14623983 T->C) n. Europeans
• • • • • • • • • •I2a2a1b2a3 Y7202 (2913976 G->A) Germans
• • • • • • • •I2a2a1b2b L623 (16202490 A->T)
• • • • • • • •I2a2a1b2c CTS11871/S3673 (23326304 C->T) Brits, Germans
• • • • • • • •I2a2a1b2d Y5188 (7838639 G->A) Maltese
• • • • • • •I2a2a1b3 S12606 (9879942 T->G)
• • • • • • • •I2a2a1b3a CTS12640/S4904/ZS21 (28652639 C->T)
• • • • •I2a2a2 S9403/SK1254 (6742730 T->C)
• • • • • •I2a2a2a L1228 (15446045 G->C) w. Europeans
• • • • • •I2a2a2b Y6099 (7527993 G->A)
• • • •I2a2b L38/S154 (15668070 A->G)
• • • • •I2a2b1 L533/S295 (2887198 G->C) n.w. Europeans
• • • • •I2a2b2 S2606 (22527402 C->A)
• • • • • •I2a2b2a S2523 (14951410 C->T)
• • • • • • •I2a2b2a1 S24121 (22201497 C->A)
• • • • • • • •I2a2b2a1a S11558 (8491250 G->A)
• • • • • • • •I2a2b2a1b F780.2 (6384078 G->A)
• • • • •I2a2b3 M8990/S3846 (7534645 A->C)
• • • • • •I2a2b3a S2488 (12733783 T->A) English
• • • • • •I2a2b3b S4556 (14124389 A->G)
• • • • •I2a2b4 P15.2/PF3112.2 (3244026 C->T)
• •I2b L415/S435 (2888663 C->T) or L417/S332 (8426321 T->C) Germans, Italians
• •I2c L596/PF6907/S292 (14197631 G->A) or L597/S333 (18887888 T->A)
• • •I2c1 PF3892 (8487200 T->C)
• • • •I2c1a L1251 (2888598 C->T) Germans
• • • •I2c1b CTS4092 (15391894 G->A)
• • • • •I2c1b1 CTS7767.1 (17651573 C->A)
• • • • • •I2c1b1a PF6328 (21797244 C->A)
• • • • • • •I2c1b1a1 F2044 (15944600 T->A) Brits
• • • •I2c1c S6685 (14185048 G->C)
• • • • •I2c1c1 S6648 (14459289 T->A)
• • • • • •I2c1c1a FGC18548 (7380574 A->G) Sardinians
• • • • • •I2c1c1b S6716 (16173282 G->C)
• • • • • • •I2c1c1b1 S6622 (14264992 A->G)
• • • • • • • •I2c1c1b1a S6595 (21430083 G->A)
• • • • • • • • •I2c1c1b1a1 S6656 (17410728 G->C)
• • • • • •I2c1c1c S9234 (6512608 A->G)
• • •I2c2 PF3827 (22444389 T->A)
• • •I2c3 S6800 (16180130 G->A)

External links

Notes and References

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Battaglia V, Fornarino S, Al-Zahery N, Olivieri A, Pala M, Myres NM, King RJ, Rootsi S, Marjanovic D, Primorac D, Hadziselimovic R, Vidovic S, Drobnic K, Durmishi N, Torroni A, Santachiara-Benerecetti AS, Underhill PA, Semino O . 6 . Y-chromosomal evidence of the cultural diffusion of agriculture in Southeast Europe . European Journal of Human Genetics . 17 . 6 . 820–830 . June 2009 . 19107149 . 2947100 . 10.1038/ejhg.2008.249 .
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Book: Underhill PA, Myres NM, Rootsi S, Chow CT, Lin AA, Otillar RP, King R, Zhivotovsky LA, Balanovsky O, Pshenichnov A, Ritchie KH . 6 . New phylogenetic relationships for Y-chromosome haplogroup I: Reappraising its Phylogeography and Prehistory . Rethinking the Human Evolution . Mellars P, Boyle K, Bar-Yosef O, Stringer C . 2007 . 33–42 . McDonald Institute for Archaeological Research . 978-1-902937-46-5 .
Mršić G, Gršković B, Vrdoljak A, Popović M, Valpotić I, Anđelinović Š, Stenzl V, Ehler E, Urban L, Lacković G, Underhill P, Primorac D . 6 . Croatian national reference Y-STR haplotype database . Molecular Biology Reports . 39 . 7 . 7727–7741 . July 2012 . 22391654 . 10.1007/s11033-012-1610-3 . . 18011987 .
Kovacevic L, Tambets K, Ilumäe AM, Kushniarevich A, Yunusbayev B, Solnik A, Bego T, Primorac D, Skaro V, Leskovac A, Jakovski Z, Drobnic K, Tolk HV, Kovacevic S, Rudan P, Metspalu E, Marjanovic D . 6 . Standing at the gateway to Europe--the genetic structure of Western balkan populations based on autosomal and haploid markers . PLOS ONE . 9 . 8 . e105090 . 2014 . 25148043 . 4141785 . 10.1371/journal.pone.0105090 . free . 2014PLoSO...9j5090K .
Fóthi E, Gonzalez A, Fehér T, Gugora A, Fóthi Á, Biró O, Keyser C . Genetic analysis of male Hungarian Conquerors: European and Asian paternal lineages of the conquering Hungarian tribes . Archaeological and Anthropological Sciences . 12 . 1 . 2020 . 10.1007/s12520-019-00996-0. free . 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..
PhD . O.M. Utevska . 2017 . Генофонд українців за різними системами генетичних маркерів: походження і місце на європейському генетичному просторі . The gene pool of Ukrainians revealed by different systems of genetic markers: the origin and statement in Europe . National Research Center for Radiation Medicine of National Academy of Sciences of Ukraine. uk. 219–226, 302.
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Šarac J, Šarić T, Havaš Auguštin D, Novokmet N, Vekarić N, Mustać M, Grahovac B, Kapović M, Nevajda B, Glasnović A, Missoni S, Rootsi S, Rudan P . 6 . 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 . 837–845 . November 2016 . 27279290 . 10.1002/ajhb.22876 . 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). . 25873634 .
Balanovsky O . Toward a consensus on SNP and STR mutation rates on the human Y-chromosome . Human Genetics . 136 . 5 . 575–590 . May 2017 . 28455625 . 10.1007/s00439-017-1805-8 . 3714493 . 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. .
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