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Haplogroup I1 (Y-DNA)
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Haplogroup I is the oldest haplogroup in Europe and in all probability the only one that originated there (apart from deep subclades of other haplogroups). It is thought to have arrived from the Middle East as haplogroup IJ sometime between 45,000 and 30,000 years ago, and developed into haplogroup I approximately 25,000 years ago. In other words, Cro-Magnons most probably belonged to IJ and I (alongside older haplogroups like F and possibly even C).
The I1 branch is estimated to have split away 20,000 years ago and evolved in isolation in Scandinavia during the late Paleolithic and Mesolithic. I1 is defined by at least 15 unique mutations, which indicates that this lineage experienced a serious population bottleneck. Men belonging to this haplogroup all descend from a single ancestor who lived between 10,000 and 7,000 years ago.
During the Mesolithic period, pre-I1 and I1 people were part of the sucessive Erteb°lle culture (5300-3950 BCE), Funnelbeaker culture (4000-2700 BCE) and Pitted Ware culture (3200-2300 BCE). The latter two are sometimes considered as Neolithic cultures due the introduction of farming. However, Neolithic farmers from Germany penetrated late into Scandinavia and in small numbers, and the lifestyle remained primarily one of hunter-gatherers. This is probably the reason why Scandinavia retained one of the most substantial Paleolithic ancestry in Europe.
The big cultural and genetic upheaval came with the Indo-Europeans from Eastern Europe who brought Scandinavia into the Bronze Age with a very short Neolithic transition. The first Indo-Europeans to reach Scandinavia were the Corded Ware (3200-1800 BCE) people from modern Russia, Belarus and Poland, who are thought to have belonged predominantly to haplogroup R1a. These people carried similar maternal lineages as Scandinavian I1 inhabitants - in great majority U4 and U5 lineages.
The second major Indo-European migration to Scandinavia was that of haplogroup R1b, the branch that is thought to have introduced Proto-Germanic languages, as an offshoot of the Proto-Celto-Germanic speakers from Central Europe. R1b probably entered Scandinavia from present-day Germany as a northward expansion of the late Unetice culture (2300-1600 BCE).
According to the Germanic substrate hypothesis, first proposed by Sigmund Feist in 1932, Proto-Germanic was a hybrid language mixing Indo-European (R1b, and to a lower extent R1a) and pre-Indo-European (native Scandinavian I1) elements. This hybridisation would have taken place during the Bronze Age and given birth to the first truly Germanic civilization, the Nordic Bronze Age (1700-500 BCE).
Subclades and Haplotypes
Grouping by STR
All Germanic tribes expanded from a small geographic core around Denmark and southern Sweden within the last 2500 years. STR variations allows to divide I1 members in various categories. There are two main clusters, each with their own subgroups.
- the Northern cluster, peaking in Norway, Sweden and Finland, which corresponds to the I1a2 (L22+, formerly known as I1d) subclade. It normally has an STR value greater than 22 for DYS390.
- the Norse group, corresponds to Ken Nordtvedt's Norse (mostly Swedish) and Ultra-Norse (mostly Norwegian and Icelandic) haplotypes. The Ultra-Norse haplotype 1 (I1-uN1) difers from the Norse one by having DYS385b=15 and (usually) DYS449=29.
- the Bothnian group, is found mostly in Finland and northeast Sweden. It corresponds to I1a2c1 (L258+) subclade, which it makes up 75% of the I1 lineages in Finland.
- the Southern cluster, most common in Denmark, Germany, the Low Countries and the British Isles. It corresponds to Ken Nordtvedt's Anglo-Saxon haplotype (originally Danish and North German).
- the Danish/Polish group usually has a DYS557 value greater than 15.
- the Western group, comprising the Low countries, England, Scotland and Ireland, matches the Z58+ subclade. It probably matches Anglo-Saxon and Frisian/Batavian ancestry.
- there appears to be a specific Welsh subgroup defined by a GATA-H4 value superior or equal to 11. This subgroup is also found in England and on the continent, but is especially common in Wales.
- the German group, is the most common type of I1 in Germany, France, Italy and Central Europe, but is also found in the British Isles and to a lower extent in Scandinavia. It is defined by a DYS456 value inferior to 15. It corresponds to the Z63+ subclade.
- Z131+ is a minor subclade that has been found in areas bordering the ancient Celtic-Germanic boundary (Belgium, central Germany, Bohemia).
- DF29+ represents the vast majority of I1 lineages.
- M227+ has so far been found only in the Baltic countries, Belarus, Poland, Switzerland, France and southwest England (Devon). It has a peculiar distribution lying at the confines of Germanic-speaking regions, although it has never been found in the core of ancient Germanic culture.
- L22+ (aka S142+) is the main Nordic subclade. It is also very common in Britain, especially on the east coast where the Vikings settled most heavily, in the Low Countries and Normandy (also probable Danish Viking heritage), as well as in Poland and Russia (Swedish Vikings).
- P109+ A mostly southern Scandinavian subclade, with a presence in all the regions settled by the Danish Vikings. It has been found sporadically in many parts of Europe, such as western Iberia, northern Italy, the Balkans, Lithuania and Russia.
- L205+ is mostly limited to the Low Countries, France and Britain. Isolated cases were also identified in Sweden and Spain.
- L287+ is an overwhelmingly Finnish subclade (found nation-wide), with a very small presence in Norway, Sweden, Poland and Russia.
- L300+ is a minor subclade found almost exclusively in southern Finland.
- L813+ is a predominantly Scandinavian subclade, particularly common in southern Norway. It is also found in Britain (likely of Viking descent) and in the northern Netherlands (but not in Germany).
- Z58+ is chiefly West Germanic, with a very strong presence in Germany, the Low Countries and Britain. It is also found to a lower extent in Nordic countries and throughout Continental Europe. Its age has been estimated around 4000 years before present.
- Z138+ (aka Z139+) is a very disparate subclade. It is found at very low frequency thoughout the Germanic world, with a peak in England and Wales (although it could just be because of oversampling in Britain). It has also been found in Portugal, southern Italy and Romania. Z138+ corresponds to AS2, AS10, AS1010.2, AS10910, AS1221 and AS1414 and Esc-13 in FTDNA's STR-based nomenclature.
- Z59+ is the main branch of Z58.
- Z382+ has been found especially in the British Isles and Germany, with a minor presence in Scandinavia. Isolated samples were also identified in the Netherlands, Finland, Italy, Croatia and Romania. corresponds to AS3, AS3-911, AS13 and Sw in FTDNA's STR-based nomenclature.
- Z60+ is spread throughout the Germanic world
- Z140+ is a strongly West Germanic subclade, found essentially in the British Isles the Low Countries, northern France, central and southern Germany, and Switzerland. It is very rare in Nordic countries. Isolated samples were found in Spain, central and southern Italy, Slovenia, Bohemia, Poland, Ukraine and Russia. Z140* matches the AS5, AS6, AS814 and EE haplotypes in FTDNA's STR-based nomenclature.
- L338+ has a similar distribution to Z140. It typically has the STR value GATA-H4=9. L338+ corresponds to AS1, AS1H, AS8, AS114 and AS11616 in FTDNA's STR-based nomenclature.
- Z73+ (and L1301+) is chiefly northern Scandinavian and Finnish. It is also found in Russia, on the east coast of Britain and the Scottish Isles (Viking heritage). Z73+ corresponds to AS9 and AS16 in FTDNA's STR-based nomenclature.
- L573+ appears mainly North German, with isolated samples in France and Sweden.
- L1248+ (and L803+) is a minor subclade that has been found in Britain, Germany and Russia.
- Z63+ is a strongly German subclade, virtually absent from Nordic countries. It is most common in Central Germany, the Benelux, England, Lowland Scotland, as well as Poland. It has also been found in Russia, Ukraine, the Balkans, Italy, Spain and Portugal.
Distribution of haplogroup I1 in Europe
Haplogroup I1 is the most common I subclade in northern Europe. It is found mostly in Scandinavia and Finland, where it typically represent over 35% of the male Y-chromosomes. Associated with the Norse ethnicity, I1 is found in all places invaded by ancient Germanic tribes and the Vikings. Other parts of Europe speaking Germanic languages come next in frequency. Germany, Austria, the Low Countries, England and the Scottish Lowlands all have between 10% and 20% of I1 lineages.
The presence of I1 is Finland can be attributed to an expansion from Sweden. The timing of this expansion is more problematic. 28% of the population of Finland is I1 (45% in Western Finland), among which 75% belong to the I1a2 subclade (mostly I1a2c1-Bothnian). I1 is accompanied by 7.5% of R1a and 3.5% of R1b. R1a is thought to have reached Scandinavia in the early Bronze Age, and R1b in the late Bronze Age. It might consequently make sense to date the first Scandinavian colonisation of Finland to the late Bronze Age. However, three elements point at an older diffusion:
Most of the Suomi I1 fits with the I1a2c and I1a2c1 subclades, two subclades that are almost exclusive to Finland. They are only found at low frequency in Sweden and Norway, and not elsewhere, despite the numerous migrations within Scandinavia from the Bronze Age onwards. If I1 arrived in Finland during the Bronze or Iron Age, Finland would have a similar diversity as Scandinavia.
Finland is the only country with more than 15% of I1 where the Germanic culture and language didn't take root. A good reason for this would be that Germanic culture didn't exist in Scandinavia at the time of the migration because the Indo-Europeans (R1a and R1b) hadn't arrived yet.
The very high frequency of I1 in Finland compared to R1a and R1b also disagrees with a Bronze Age dispersal. It is undeniable that the Corded Ware culture, associated with the first diffusion of R1a, expanded as far as Finland. Half of the R1a in Finland matches the Polish cluster (associated with the Corded Ware), while the other half seems to have come later from Scandinavia. It is likely that the Corded Ware expanded by the Baltic countries to reach Finland, rather than go all the way to Germany, Denmark, Sweden then Finland. This first wave of R1a to Finland would therefore not have brought any I1 with it. The diffusion pattern of R1a from that the Corded Ware period show that R1a people advanced without assimilating indigenous people on the way (otherwise there would probably be more I2a and E1b1b in Scandinavia). So there is no reason to believe that a minority of R1a led a population eight times as big of I1 men from Scandinavia to Finland. Most of the Finnish R1b belongs to the Germanic R1b-S21 subclade. In other words, the Finns have roughly 3% of R1a and 3% of R1b of Germanic/Scandinavian origin, so in all likelihood no more than 3 or 4% of I1 from that period as well. The bulk of I1 (most of them I1d), approximately 25% of the Finnish male lineages, must have reached Finland before the Bronze Age, or possibly as a displacement of part of the indigenous Scandinavian I1 population by the Bronze-age invaders.
Unfortunately this timeline seriously conflicts with the estimated age of I1a2c, which Ken Nordtvedt calculated to be only 2000 years old based on STR variatons. This method is not very accurate because it fails to take into account population size. Larger populations create more genetic variations. Nordic countries have always had a lower population density than central of southern Europe. Before the Bronze Age, Nordic people were still hunter-gatherers, while the rest of Europe had been farming for up to 3500 years. Agricultural societies could support populations ten times higher than hunter-gatherers in similar climates. In cold Fennoscandia, the pre-Indo-European population density must have been at least 20 times lower than in Mediterranean Europe. This means that the mutation rate would also be 20 times lower, and therefore that haplogroup I1 is much older than STR variations alone would suggest. If the age estimate of 2000 years old happened to be correct anyway (very unlikely), the only way I1 could have become so predominant in Finland is through an unprecedented founder effect, with a single male lineage quickly replacing one fourth of all lineages in the country.
I1 in the rest of Europe
The Germanic migrations dispersed I1 lineages to Britain (Anglo-Saxons), Belgium (Franks, Saxons), France (Franks, Visigoths and Burgundians), South Germany (Franks, Alamanni, Suebi, Marcomanni, Thuringii and others), Switzerland (Alamanni, Suebi, Burgundians), Iberia (Visigoths, Suebi and Vandals), Italy (Goths, Vandals, Lombards), Austria and Slovenia (Ostrogoths, Lombards), Ukraine and Moldova (Goths), as well as around Hungary and northern Serbia (Gepids). The I1 found among the Poles (6%), Czechs (11%), Slovaks (6%) and Hungarians (8%) is also the result of centuries of influence from their German and Austrian neighbours. An interesting hotspot is Macedonia (10%), which could be the descendants of the Goths who settled in the Eastern Roman Empire in the 3rd and 4th centuries.
The Danish and Norwegian Vikings brought more I1 to Britain, Ireland, the Isle of Man, Normandy, Flanders, Iberia, Sicily... The Swedish Vikings (Varangians) set up colonies in Russia and Ukraine, and outposts as far as the Byzantine Empire, the Caucasus and Persia. The higher frequency of I1 in Northwest Russia (east of the Baltic) hints at had a particularly strong Varangian presence, which is concordant with the establishment of the Kievan Rus' by the Swedes.
|Malmstr÷ma et al. tested the DNA of Birger Jarl and his son Eric Birgersson, and they were identified as members of haplogroup I1. Birger Jarl was one of Sweden's greatest medieval statesman. He was the founder of Stockholm, and acted as regent of Sweden for 18 years. His sons Valdemar and Magnus suceeded each others as kings of Sweden, and their descendants for one hundred years. This dynasty is known as the House of Bjelbo, and all six kings were presumably members of haplogroup I1.|
|The Hamilton DNA Project compared the Y-DNA of four descendants of Alexander Hamilton, one of the a Founding Fathers of the United States of America. All shared very close STR values, proving beyond reasonable doubt that they shared a same recent patrilineal ancestor.|
|The haplogroup of the celebrated Russian writer Leo Tolstoy was inferred by testing one of his descendants, Pyotr Tolstoy.|
|The personal genomics company 23andMe tested the Y-chromosomal haplogroup of business magnate and multi-billionaire Warren Buffett and singer/film producer Jimmy Buffett to determine whether they shared a common patrilineal ancestor. Both men were found to belong to haplogroup I1, though apparently not to closely related subclades.|