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суббота, 10 января 2015 г.

Ancient Ancestry Project: Benelux & France

Y-chromosomal DNA Project for France, Belgium, Luxembourg, and the Netherlands

Last update January 2015


A genetic history of France and the Low Countries


Paleolithic & Mesolithic hunter-gatherers

The history of Western Europeans starts approximately 35,000 years ago with Cro-Magnons, the first Homo Sapiens to colonise Europe. They left the Near East 45,000 years ago and progressed very slowly from southeastern to western Europe, probably intermingling with the autochthonous Neanderthals on the way. Only one Paleolithic Y-DNA has beentested (by Seguin-Orlando et al. 2014 to date, a 37,000 year-old individual from southern Russia, and belonged to Y-haplogroup C1 and mt-haplogroup U2. Haplogroup C1 (actually C1a2) also shows up in a sample from Mesolithic Spain 7,000 years ago (Sánchez-Quinto et al. 2012) - a surprising continuity in both time and space.


It likely that the first migrations of Homo sapiens to Europe, linked with the development of Aurignacian culture, brought old haplogroups such as C-V20 and F-P96, simply because other haplogroups further down the phylogenetic tree did not exist yet. It is possible that even the West African haplogroup A1a reached Europe during the Paleolithic by crossing the Strait of Gibraltar, as some sub-Saharan African admixture regularly shows up in Mesolithic and Paleolithic samples, and modern A1a samples has been found in the northern fringes of Europe (Ireland, Norwegian Lappland, Finland) which cannot be explained by any more recent migration.
The robust people of the Aurignacian were succeeded by more gracile people during the Gravettian period (32,000-22,000 years ago). This second wave of modern humans is probably the one that brought haplogroup I. A third wave in the late glacial period (17,000-12,000 years ago) might have brought E-V13 from North Africa to Iberia, Italy and Greece, while J2b expanded from Anatolia to the southern Balkans.

During the Mesolithic period (c. 12,000-7,000 years ago), starting at the end of the last Ice Age around 10,000 BCE and lasting until the adoption of agriculture (from circa 5500 BCE in southern France until 4000 BCE in the northern Netherlands), the inhabitants of the Benelux and France would have belonged primarily to haplogroup I2, and particularly to I2a1a (like the 8,000 year-old Loschbour man tested by Lazaridis et al. 2014), although I2a2 might have been present between Alsace and Franche-Comté and the Netherlands. E1b1b might have by then expanded from Iberia and Italy into southern France.

Neolithic & Chalcolithic herders, farmers and megalith builders

The Neolithic period saw the advent of agriculture (livestock herding, then cereal farming), the use of pottery and the replacement of the nomadic lifestyle by sedentism. Originating in the Fertile Crescent circa 9500 BCE, the Neolithic spread through Anatolia and Greece, then its diffusion split in two routes. The southern route went across the Mediterranean to Italy, southern France and Iberia, and evolved into the Printed-Cardium Pottery culture (5000-1500 BCE). The northern branch developed in the Balkans as the Starčevo culture (6000-4500 BCE) and the Linear Pottery culture (LBK) (5500-4500 BCE), following the Danube and its tributaries until Germany, northern France and the Benelux. It has been suggested that the modern divide between northern European cultures, favouring butter-based cooking, and southern European cultures, preferring olive oil-based cuisine, dates back from the Neolithic period.

 

Neolithic herders and farmers from the Near East appear to have belonged predominantly to haplogroup G2a, although it cannot be excluded that some herders also belonged to J1 and T, especially around the Mediterranean basin. Ancient DNA tests have confirmed the present of G2a in all Neolithic sites in Europe tested to date, including numerous samples from the Cardium Pottery, Starčevo and and Linear Pottery cultures. Isolated samples of haplogroups E-V13, F, I1 and I2 have all been found on different Neolithic sites next to a G2a majority and probably represent assimilated hunter-gatherers.
In Southwest Europe, the Cardium Pottery culture gave way to the Megalithic culture (5000-2000 BCE), which spread all over the Atlantic region, with particularly strong presences in Portugal, Galicia, Brittany and the British Isles. Megalithic people were very probably a blend of native I2 and E-V13 people and Near Eastern immigrants (G2a, and probably also J1 and T in Iberia and Italy). During the late Megalithic period, the Neolithic society progressively evolved into a Chalcolithic (Copper Age) one, apparently without significant societal changes, nor major external gene flow. The great upheaval in both culture and genes came during the Bronze Age, with the arrival of the Indo-European speakers, who originated with the Yamna culture (3500-2300 BCE) in the Pontic Steppe, to the north of the Black Sea.
Maternal lineages brought by Neolithic farmers from the Balkans and Anatolia can be safely ascertained by the large number of ancient mtDNA already tested. They included haplogroups HV, H2a2, H5a, H13, H20, J1c, K1a, N1a, T2 and X.

Bronze & Iron Age Indo-Europeans

There were two distinct, though related, genetic groups of Proto-Indo-Europeans: the southern R1b branch, linked with the diffusion of the Greco-Anatolian, Albanian, Italic, Celtic and Germanic languages, and the northern R1a branch, associated with the propagation of the Daco-Thracian, Illyrian, Balto-Slavic and Indo-Iranian languages.
R1a was the first to reach the Low Countries and northeastern France, with the Corded Ware culture (2900-2400 BCE), a late Chalcolithic and early Bronze Age expansion of the Yamna culture. Lineages dating from this period would belong to the Z283 subclade.
The Proto-Celto-Germanic branch of R1b (L11) settled around Bohemia and eastern Germany circa 2800 BCE and established the Unetice culture, the bronze-age culure which would expand across all Western Europe and Scandinavia over the next millennium, and replace the Neolithic/Chalcolithic Bell Beaker culture.

An expansion of Unetice to the north and west gave birth to the Proto-Germanic branch (R1b-U106), which mixed with the indigenous populations of northern Germany and the Netherlands, notably I2a2 (descended from Cro-Magnons) and R1a-Z283 (from the Corded Ware culture), but also with a minority of Neolithic lineages (E-V13, G2a, J, T). From 1700 BCE, R1b-U106 people penetrated into Scandinavia, where they blended with the local I1 and R1a-Z283 populations.
Judging from the propagation of bronze working to Western Europe, those first Proto-Celts reached France and the Low Countries by 2200 BCE, then the British Isles by 2100-2000 BCE. This first migration would have brought the L21 subclade of R1b to Northwest Europe. Through a founder effect, L21 became the dominant paternal lineage among the ancient Britons and Irish, and remained it among modern Bretons, Welsh, Highland Scots and Irish. Another migration from Germany appears to have been led by men belonging to the DF27 subclade of R1b and conquered Southwest France, then the Iberian peninsula. DF27 is now by far the main paternal lineage of the Gascons, Basques and Catalans.

The third major Proto-Celtic branch was R1b-U152, which is thought to have evolved from the Urnfield, Hallstatt and La Tène cultures (1300-50 BCE) around the Alps. An early Urnfield migration brought U152 to Italy around 1200 BCE, where they became Italic speakers, including the Romans (=> see Genetic history of Italy). The La Tène culture (450-50 BCE) is the one most strongly associated with the ancient Gauls. Gaul encompassed all modern France, and all the Low Countries south of the Rhine, as well as the German Rhineland, which happens to be the territory where U152 is found at the highest frequencies outside Italy. U152 can therefore be considered to be a marker of both Gaulish and Italic ancestry. U152 is divided in many subclades of its own, some of which are Italic, while others are Gaulish or more widely associated with the Hallstatt and La Tène expansions across a vast part of Europe, as far west as Iberia and England, and as far east as Anatolia and Ukraine or Russia (and perhaps even China). The Gaulish invasions of northern Italy, the latter Roman conquest of Gaul, and the numerous intermarriages across the two sides of the Alps mean that both Italic and Gaulish subclades are found scattered across Western Europe nowadays. However, the L2 and Z36 subclades appear to be mostly northern and therefore probably more Celtic, while Z144 and Z192 subclades are far more frequent in Italy and could be more genuinely Italic or Roman.

Classical Antiquity & Middle Ages

Between 600 BCE and 300 BCE the Greeks founded colonies along the Mediterranean coast of France, founding Agde, Marseille, Hyères and Nice. The ancient Greeks would have brought chiefly haplogroups E1b1b and J2 with them, with also a minority of G2a, J1, R1b-L23 and T. Then came the Romans, who stayed for 500 years in Gaul. The Romans are thought have belonged essentially to R1b-U152, with substantial minorities of E1b1b (probably more E-M123), G2a (especially G2a3b1a), J1, J2 (both J2a and J2b2), and T.
During the late Roman period, the Franks were allowed to settle peacefully within the borders of the empire around the territory of modern Belgium. The Franks were a Germanic tribe who originated somewhere between the northern Netherlands and Denmark. They would have belonged mostly to haplogroups R1b-U106 (about half of all lineages), I1, R1a (Z283 and L664) and I2a2a (M223).

In the 5th century, the Burgunds, the Visigoths and other Germanic tribes invaded Gaul under of the pressure from the Huns, a tribe that originated around modern Mongolia. The Burgunds originally came from the island of Borgholm in eastern Denmark, while the Goths hailed from southern Sweden. Both tribes would have carried a significantly higher percentage of haplogroups I1 and R1a than the Franks. The Goths had lived for several centuries in Poland, Ukraine and the Balkans before reaching southern France and could have been carrying Proto-Slavic R1a (Z280 and M458) as well as Balkanic lineages (I2-M423, E-V13, J2b). The genetic influence of the Goths in France and Iberia appears to have been very minor though. The Huns, who would have belonged to haplogroups C3, Q1a and R1a-Z93, may account for a tiny fraction of modern paternal lineages in France and Belgium.
After the fall of the Western Roman Empire, the Franks re-unified ancient Gaul under their rule and conquered most of Germany, Austria, Switzerland and the northern half of Italy (=> see History of the Franks).
Between the 9th and the 11th centuries the Vikings settled in various regions of France and the Benelux (notably Normandy and Bruges), bringing a new influx of Germanic haplogroups (R1b-U106, R1a-Z283, I1, I2-M223), but this time with a higher proportion of I1, especially the Nordic I1a2 (L22) subclade, but also the typically Scandinavian R1a-Z284 subclade.

Benelux & France Y-DNA Project

What was the genetic impact of these invaders on the present population of these countries ? Where did the Romans settle most heavily ? Are there pockets of isolated ethnicities in unexpected parts of France ? All these questions can be answered with your help.
Discovering the genetic rift between northern and southern Germany is one of the achievements of genetics that has given us a new look on what defines ethnicities in Europe. It has proven that ethnicities do not match modern linguistic groups. France looks even more interesting due to its size, varied relief with pockets of isolated communities and minority languages.
What about Belgium, this small country divided by unending linguistic quarrels ? Do Flemings and Walloons really have different origins ? It is easy to claim that the Flemings are of Germanic descent just because they speak a Germanic language. But aren't Walloons also of Germanic descent ? History tells us that the Franks settled mostly along the Meuse valley and in the Hainaut, so in northern Wallonia. What will DNA say ? Did the ancient Belgae survive the Roman and Frankish conquests ? If so, what percentage of the population can claim Celtic ancestry ?

The purpose of this project is to estimate the percentage of Germanic, Italo-Celtic, Near-Eastern or other ancestry in each region, province or departement of France. Luxembourg, Belgium and the Netherlands. Anyone can contribute by testing their Y-DNA with one of the dozens of commercial company.

What is a geographic DNA project ?

A geographic DNA projects aim at better understanding a region's genetic make-up and genetic history by looking at the distribution of paternal lineages inherited through the Y chromosome, known as Y-chromosomal haplogroups.

Many geographic DNA project already for most European countries. Data for France and Wallonia is particularly scarce though, which is why Eupedia thought of creating this project. DNA tests have become cheap enough in the last few years for almost anyone to participate in such a project. Here are the tests that we recommend, with the pros and cons for each.

  • 23andMe: for only 99$ you will know both your paternal and maternal haplogroups. This tests includes autosomal data which can be used to determine your admixtures on all chromosomes. It also also a report for physical and psychological traits on risks for medical conditions
  • Geno 2.0: official test of The Genographic Project by National Geographic in association with Family Tree DNA. It is similar to the 23andMe test, but is more expensive (200$) and does not provide any medical or phenotypic report. The main advantage of this test is that it is the most accurate to date for Y-DNA and the results can be used for DNA Projects and by research groups at FTDNA which aim to identify new subclades.
  • BritainsDNA: the Chromo2 complete (389$) is similar to the 23andMe test with more detailed Y-DNA, while the Chromo2 Y-DNA Raw (199$) is equivalent to the Y-DNA data from the Geno 2.0 test.
  • Family Tree DNA: The STR tests of FTDNA are mostly useful to compare individuals in genealogical times. You can determine your paternal haplogroup with their Y-DNA12 for only 49$, but it is extremely basic and you will need (costly) upgrades to determine your deep subclades. The Y-DNA67 (269$) is usually enough to determine the deep subclade, but may require additional SNP tests. The strong suit of FTDNA is their DNA Projects and research groups. But it is usually better to purchase directly the Geno 2.0 and join a project.
  • Brabant Y-DNA Project: if your paternal ancestors are from the Benelux you can join this project for 140€. The test consists of 38 STR markers + a deep subclade SNP analysis.
N.B. You can submit your results using any of these tests to join the Benelux & France DNA Project on Eupedia.

Genetic make-up of the Benelux

Based on the present data, the haplogroup distribution for the Netherlands (Holland), Belgium and Luxembourg is as follow.

Haplogroup I1 I2*+I2a I2b R1a R1b G J2 J1 E1b T L Q
Netherlands 16.5 1 6.5 4 49 4.5 3.5 0.5 3.5 1 0 0
Flanders 13 2.5 4 4 60 4 4 1 5.5 0.5 0.5 0.5
Wallonia 11 2.5 6 4 59 4 1 0 8.5 1.5 0 0
Overall 60% of Belgian men belong to R1b, including 32.3% to the Italo-Celto-Germanic P312 (S116) subclade, and 25% to the Germanic U106 (S21) subclade. Here is breakdown of R1b subclades in Belgium from the Brabant Y-DNA Project.

  • P25 : n=1 (0.1%)
    • P297 : n=1 (0.1%)
      • M269 (+ L23) : n=16 (1.8%)
        • P310/L11 : n=6 (0.7%)
          • U106/S21* : n=15 (1.7%)
            • Z18 : n=18 (2%)
            • Z381 : n=72 (8.3%)
              • U198 : n=9 (1%)
              • L48 : n=104 (11.9%)
          • P312/S116* : n=84 (9.6%)
            • L21 : n=78 (8.9%)
            • Z196* : n=27 (3.1%)
              • SRY2627 : n=6 (0.7%)
            • U152* : n=28 (3.2%)
              • L2 : n=46 (5.2%)
                • L20 : n=13 (1.5%)

Analysis

The dominant haplogroup in the Benelux is R1b, almost equally divided between the subclades R1b-U106 and R1b-P312. Both are present in all the Benelux, although R1b-U106 (S21) reaches its maximum frequency in Frisia (42%) and the central Netherlands (35%) and decreases progressively towards the south to 30% in the southern Netherlands, 25% in Flanders and 22% in Wallonia. R1b-P312 (S116) displays roughly the opposite pattern peaking in the south-east of Wallonia and Luxembourg (50%), then lowering to 32% in Flanders, 20% in the southern Netherlands, 15% in the central Netherlands and 10% in Frisia.
The Atlantic Celtic R1b-L21 (S145) lineage, most commonly found in the British Isles, reaches its maximum in the western half of Belgium (10%), including Flemish and Walloon Brabant, then decreases to 7-8% to the east of the country. Its frequency falls to 3-5% in the Netherlands, with little difference nationwide.
The Gaulish Celtic and Italo-Roman lineage R1b-U152 (S28) has the clearest north-south gradient of any haplogroup in the Low Countries. It is most common in southeastern Belgium and Luxembourg, where it can make up over 20% of the paternal lineages, then decreases to an average of 10% in all Flanders, 6% in the southern Netherlands, 3.5% in the central Netherlands, and only 1% in Frisia.
Haplogroup I1, one of the most reliably Germanic lineages, has almos identical frequencies in Flanders (13%) and Wallonia (11%), but is slightly higher in the Netherlands (16.5%), although that is still a far cry from the 35% observed in Scandinavia.
Near Eastern haplogroups (E1b1b, G2a, J1, J2, T) , which were spread mostly by Neolithic herders and farmers, then to a much lower extent also by the Romans, make up approximately 15% of the population, with little difference between the regions. There seems to be an inverted north-south gradient in the proportions of J1-J2 and E1b1b-T, although it is not clear why. Haplogroups J1 and J2 are the two most common Jewish lineages. J1 was only found around Amsterdam and Antwerp, two cities known for welcoming Jewish immigrants in past centuries, while J2 was also higher in both in Holland and in the province of Antwerp. It is therefore likely that the differential of 2% for these lineages in Holland and Antwerp are of Jewish origin.
The more elevated frequencies of E1b1b and T in Wallonia is probably a remnant from the Early Neolithic. Most of the E1b1b is E-V13, the most common subclade in Central and Eastern Europe, linked to the diffusion of the Linear Pottery Culture. The percentage of E1b1b and T in Wallonia is equivalent to the one in western and southern Germany. Wallonia also shares a particularly high percentage of I2-M223 with western and central Germany, hinting that the Walloons may be closer in some respect to central-west Germans, while the Flemish and the Dutch resemble more north-west Germans. The main difference is that both the Dutch and the Belgians have considerably lower levels of haplogroup R1a than all the Germans.
The sample size for Wallonia is too small (n=81) to draw conclusions about inter-provincial variations, but the present data indicates that the west (Hainaut) has a very high percentage of R1b-U106, while the east and south (Liège and Luxembourg) are overwhelmingly R1b-U152, which is either of La Tène/Gaulish Celtic or Roman origin. The province of Namur in central Wallonia (Meuse valley) has a surprisingly high percentage of Germanic haplogroups - over 50% for I1, I2a2a, R1a and R1b-U106 combined.

Genetic make-up of France

Here is the haplogroup distribution in France based on the data collected so far. Click on a haplogroup name to sort the table by frequency. Over 5000 samples were taken into account. Note that the vast majority (> 80%) of samples could not be linked to a particular region of France because the source study did not only mentioned the region, or, in the case of commercial samples, the individuals tested did not know for sure region of patrilineal ancestry, as is common for North Americans and Parisians. Only half of the French regions had at least 100 samples. The others could not be listed here as the frequencies would be completely meaningless. The large number of unsorted samples explains the discrepency between the total of the regional data (including mostly outlying regions like Brittany, Flanders-Artois, Alsace, Gascony and Corsica) and the total for France (which includes probably much more data from central regions).

Region/Haplogroup I1 I2*+I2a I2b R1a R1b G J2 J1 E1b T Q
Alsace 8 1 3 3 55 6 8 1 10 4 1
Auvergne 2 1 1.5 5.5 52.5 9 8 3.5 12.5 4.5 0
Brittany 8 1 4.5 0.5 80 2 2.5 0.5 0.5 0 0
Corsica 0 18.5 1 0 49 7.5 14 0 8 0.5 0
Flanders-Artois 13 2.5 1 2.5 61 6 2.5 0 12 0 0
Gascony 0 4.5 0 0 91 0 4.5 0 0 0 0
Ile-de-France 2.5 2 2 3.5 57.5 3.5 6.5 1 20.5 0 0
Provence 2 4 1 5 58 7.5 8.5 1 10.5 1 2
Midi-Pyrénées 2.5 5 2.5 4 61 4 8.5 4 7.5 1 0
Normandy 7 4 2 1 76 0 2 1 5 0 1
Poitou-Saintonge 1.5 3 0 0 74.5 7.5 6 0 1.5 0 0
Rhône-Alpes 7 1.5 2.5 5 66.5 5 2.5 0 5 0 5
TOTAL FRANCE 8.5 3 3.5 3 58.5 5.5 6 1.5 7.5 1 0.5

Analysis

Note that the total for France is biased towards North Americans of French descent (mostly from Québec), as genealogical DNA tests have not yet become popular among French people.
R1b is the most common haplogroup in France. It includes four main subclades : the Atlantic Celtic R1b-L21 in the north-west, the Gascon-Iberian R1b-DF27 (including the Basque R1b-M153) in the south-west, the Germanic R1b-U106 in the north, and the Gaulish Celtic and Italic R1b-U152 in the east.
The ancient Burgundians, a Germanic tribe from eastern Denmark, appear to have carried considerable percentages of haplogroups R1a and Q, two haplogroups that are now found at unusually high frequencies around the former Kingdom of the Burgundians, in what is now the Rhône-Alpes region and the north of Provence.

Geographic distribution of ethnic features in France

This map shows an estimation of the dominant ancestry in each region of France based on anthropological studies. Will DNA confirm this general pattern ? Here is a summary of Y-DNA haplogroups found in France, and the ancient ethnicities associated with them :

  • Germanic/Nordic : R1b-U106, I1, I2-M223
  • Gaulish Celtic : R1b-U152, R1b-L21, R1b-P312
  • Atlantic Celts : R1b-L21
  • Iberian Celts : R1b-DF27
  • Basque : R1b-DF27, R1b-M153
  • Roman : R1b-U152, R1b-L23, J2, G2a, E-M123, T
  • Greek : E1b1b, J2, R1b-L23, G2a, T, J1





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