The information about the origin and ethnic association of haplogroups on this website should not be read as hard facts, but, as is often the case in science, as a model in constant evolution based on the present knowledge and understanding (of the authors). Whenever the advancement of genetics couldn't provide irrefutable answers, we have attempted to provide the most likely and logical hypothesis based on archeological, historical and linguistic evidence. This page is being updated regularly to keep up with recent studies giving additional insights or rectifying possibly erroneous theories. Feel free to add comments or share your opinion on the forum.
Introduction to genetic genealogy
DNA studies have permitted to categorise all humans on Earth in genealogical groups sharing one common ancestor at one given point in prehistory. They are called haplogroups. There are two kinds of haplogroups: the paternally inherited Y-chromosome DNA (Y-DNA) haplogroups, and the maternally inherited mitochondrial DNA (mtDNA) haplogroups. They respectively indicate the agnatic (or patrilineal) and cognatic (or matrilineal) ancestry.
Y-DNA haplogroups are useful to determine whether two apparently unrelated individuals sharing the same surname do indeed descend from a common ancestor in a not too distant past (3 to 20 generations). This is achieved by comparing the haplotypes through the STR markers. Deep SNP testing allows to go back much farther in time, and to identify the ancient ethnic group to which one's ancestors belonged (e.g. Celtic, Germanic, Slavic, Greco-Roman, Basque, Iberian, Phoenician, Jewish, etc.).
In Europe, mtDNA haplogroups are quite evenly spread over the continent, and therefore cannot be associated easily with ancient ethnicities. However, they can sometimes reveal some potential medical conditions (see diseases associated with mtDNA mutations). Some mtDNA subclades are associated with Jewish ancestry, notably K1a1b1a, K1a9,d K2a2a and N1b.
DNA Facts
Nucleotides are the alphabet of DNA. There are four of them : adenine (A), thymine (T), guanine (G) and cytosine (C). They always go by pairs, A with T, and G with C. Such pairs are called "base pairs".
The 46 chromosomes of human DNA are composed of a total of 3,000 million base pairs.
The Y-chromosome possess 60 million base pairs, against 153 million for the X chromosome.
Mitochondrial DNA is found outside the cell's nucleus, and therefore outside of the chromosomes. It consists only of 16,569 base pairs.
A SNP (single nucleotide polymorphism) is a mutation in a single base pair. At present, only a few hundreds SNP's define all the human haplogroups for mtDNA or Y-DNA.
Y-DNA Haplogroups
Chronological development of Y-DNA haplogroups
K => 40,000 years ago (probably arose in northern Iran)
T => 30,000 years ago (around the Red Sea)
J => 30,000 years ago (in the Middle East)
R => 28,000 years ago (in the Central Asia)
E1b1b => 26,000 years ago (in southern Africa)
I => 25,000 years ago (in the Balkans)
R1a => 21,000 years ago (in southern Central Asia)
R1b => 20,000 years ago (in the Northwest Asia)
E-M78 => 18,000 years ago (in north-eastern Africa)
G => 17,000 years ago (between India and the Caucasus)
I2 => 17,000 years ago (in the Balkans)
J2 => 15,000 years ago (in northern Mesopotamia)
I2b => 13,000 years ago (in Central Europe)
R1a1 => 12,000 years ago (north of the Black Sea)
N1c1 => 12,000 years ago (in Siberia)
I2a => 11,000 years ago (in the Balkans)
R1b1b2 => 10,000 years ago (north or south of the Caucasus)
J1 => 10,000 years ago (in the Arabian peninsula)
E-V13 => 10,000 years ago (in the Balkans)
I2b1 => 9,000 years ago (in Germany)
I2a1 => 8,000 years ago (in Sardinia)
I2a2 => 7,500 years ago (in the Dinaric Alps)
E-M81 => 5,500 years ago (in the Maghreb)
I1 => 5,000 years ago (in Scandinavia)
R1b-L21 => 4,000 years ago (in Central or Eastern Europe)
R1b-S28 => 3,500 years ago (around the Alps)
R1b-S21 => 3,000 years ago (in Frisia or Central Europe)
I2b1a => less than 3,000 years ago (in Britain)
Map of early Bronze Age cultures in Europe around 4,500 to 5,000 years ago
Hypothetical map of Y-DNA haplogroup distribution in Europe about 2,000 years ago
This map was composed by calculating modern regional densities and withdrawing the supposed influence of migrations that took place in the last 2000 years. Only the main/dominant haplogroups are represented for each region. Haplogroup E and R1b encompass various subclades if the subclade not specified.
Large font = over 25% of the population Small font = between 10 and 25% of the population
European haplogroups
Haplogroup R1b (Y-DNA)
R1b is the most common haplogroup in Western Europe, reaching over 90% of the population in some parts of western France, northern Spain or Ireland. It is widespread as far as Central Asia and Africa.
Haplogroup R1* might have originated in southern Central Asia (between the Caspian and the Hindu Kush), then developed into R1b* then R1b1* in the northern part of the Middle East during the Ice Age. It presumptively moved to northern Anatolia and across the Caucasus during the early Neolithic, where it became R1b1b. The Near Eastern leftovers evolved into R1b1a (M18), now found at low frequencies among the Lebanese and the Druze.The Phoenicians (who came from modern day Lebanon) spread this R1b1a and R1b1* to their colonies, notably Sardinia and the Maghreb.
The Proto-Indo-Europeans belonged both R1a and R1b. Their homeland was in the Pontic-Caspian steppe, in what is known as the Kurgan culture (7000-2200 BCE). The presence of R1b in modern times between the Black Sea and the Caucasus hints at the Maykop culture (3500-2500 BCE) as their most plausible homeland, while the Eurasian steppes to the north were R1a territory.
R1b1b2 probably appeared during Maykop culture. It was an advanced Neolithic culture of farmers and herders, and one of the very first to develop metalworking, and therefore metal weapons. Stuck between two seas and the Caucasus, they imaginably traded actively around the Black Sea, notably with the other R1b people from northern Anatolia (those that didn't cross the Caucasus and might be the ancestors of the Hittites).
Horse were first domesticated in the neighbouring Yamna culture (modern Ukraine) approximately 3500 BCE, and chariots were invented in the north-eastern steppes around 2000 BCE. Cavalry and chariots played an vital role in the subsequent Indo-European migrations, allowing them to move quickly and defeat easily anybody they encountered. Combined with advanced bronze weapons and their sea-based culture, R1b people are excellent candidates for being the mysterious Sea Peoples, who raided the eastern shores of the Mediterranean during the second millennium BCE.
The European branch
The Indo-Europeans' bronze weapons and horses would have given them a tremendous advantage over the autochthonous inhabitants of Europe, namely the native haplogroup I (descendant of Cro-Magnon), and the early Neolithic herders and farmers (G2a and E-V13). This allowed R1a and R1b to quickly replace (in all likelihood through warfare) most of the native male lineages, although female lineages seem to have been less affected.
A comparison with the Indo-Iranian invasion of South Asia shows that 40% of the male linages of northern India are R1a, but only 20% of the female lineages could be of Indo-European origin (H, J, K, T, U). The impact of the Indo-Europeans was more severe in Europe because European society 4,000 years ago was less developed in terms of agriculture, technology (no bronze weapons) and population density than that of the Indus Valley civilization. This is particularly true of the native Western European cultures where farming arrived much later than in the Balkans or central Europe. Greece was the most advanced of European societies and was the least affected in terms of haplogroup replacement. Native European Y-DNA haplogroups (I1, I2a, I2b) also survived better in regions that were more difficult to reach or less hospitable, like Scandinavia, Brittany, Sardinia or the Dinaric Alps.
R1b1b2 is thought to have arrived in central and western Europe around 2300 BCE, by going up the Danube from the Black Sea coast. This correspond to an archeological vacuum in the old Maykop homeland, so the migration must have been on a massive scale, maybe due to pressure from other (R1a) Indo-European people from the north. There might have been several consecutive waves across the Black Sea to the Danube, but the largest one between 2500 BCE (end of the Maykop culture) and 2300 BCE (beginning of the Unetice culture).
It is doubtful that the Beaker culture (2800-1900 BCE) was already Indo-European (although they were influenced by the Corded Ware culture), because they were the continuity of the native Megalithic cultures. Nevertheless, it is undeniable that the following Unetice (2300-1600 BCE), Tumulus (1600-1200 BCE), Urnfield (1300-1200 BCE) and Hallstatt (1200-750) cultures were linked to the spread of R1b to Europe, as they abruptly introduce new technologies and a radically different lifestyle.
These Proto-Italo-Celto-Germanic R1b people had settled around the Alps by 2300 BCE, and judging from the spread of bronze working, reached Iberia by 2250 BCE, Britain by 2100 BCE and Ireland by 2000 BCE. This is assumably when the R1b-L21 lineage came to the British Isles, from southern Germany. A second R1b expansion took place from the Urnfield/Hallstatt culture around 1200 BCE, pushing west to the Atlantic, north to Scandinavia, and as far east as Greece and Anatolia (=> see Dorian invasion below).
The new Bronze Age culture flourished around the Alps (Unetice to early Hallstatt) thanks to the abundance of metal in the region, and laid the foundation for the classical Celtic culture. The Celtic Iron Age (late Halstatt, from 800 BCE) may have been brought by a more recent wave of immigrants from new Koban culture (1100-400 BCE) in the eastern Black Sea homeland.
Alpine Celts of Hallstatt are associated with the S28 (a.k.a. U152) mutation, although not exclusively. The Italic branch (also S28/U152) is thought to have entered Italy by 1200 BCE, but there were certainly several succesive waves, as attested by the later arrival of the Cisalpine Celts. The Belgae were another S28/U152 branch, an extension of the La Tène culture northward, following the Rhine, Moselle and Meuse rivers.
R1b-S21 (a.k.a. U106) is found at high concentrations in the Netherlands and northern Germany. Its presence in other parts of Europe can be attributed to the 5th- and 6th-century Germanic migrations. The Frisians and Saxons spread this haplogroup to the British Isles, the Franks to Belgium and France, and the Lombards to Austria and northern Italy. The high concentration of S21/U106 around Austria hints that it could have originated there in the Hallstatt period, or originated around the Black Sea and moved there during the Hallstatt period. In fact, southern Germany and Austria taken together have the highest diversity of R1b in Europe. Besides S21, the three major first level subclades of R1b1b2a1b (L21, S28, M167) are found in this area at reasonable frequencies to envisage a spread from the Unetice to Hallstatt homeland to the rest of western Europe.
The Hittites (2000-1200 BCE) were the first Indo-Europeans to defy (and defeat) the mighty Mesopotamian and Egyptian empires. The Hittite ruling class was plausibly an offshoot of the Maykop culture that had entered northern Anatolia via the sea route and conquered the Hattian kingdom. The Hattians might have had some R1b from the old Anatolian branch (from the early Neolithic) mixed with the other Anatolian E-M78, G2a and J2 people.
Other waves of (seaborne) R1b1b2 invaders from the Pontic-Caspian homeland are thought to have settled in Anatolia a few centuries later, where they became the Luwians, Lycians and Lydians (1450 BCE). Troy was most probably a colony to secure the trade routes of the Sea Peoples between the Black Sea and the Aegean. The Trojans were Luwian speakers related to the Hittites, with proven cultural ties to the culture of the Pontic-Caspian steppe. The first city of Troy dates back to 3000 BCE, right in the middle of the Maykop period, and exatly at the time the first galleys were made.
The Maykop culture was succeeded by the Srubna culture (1600-1200 BCE), then the Colchian culture (1200-600 BCE), which extended into the western Caucasus. Its further expansion to the south of the Caucasus correspond to the first historical mentions of the Proto-Armenian branch of Indo-European languages (around 1200 BCE).
The presence of R1b in Greece could be attributed to the Dorian invasion (1200 BCE), which correspond to the expansion of the Urnfield culture throughout Europe and Anatolia, and to the destruction of the Near-Eastern civilizations by the Sea Peoples. Greek R1b (including southern Italy) is divided between the Proto-Celtic S116/P312 and the eastern variety (known as ht35) from Anatolia. If the Dorian were ht35, they could be the descendants of the Trojans (seeking revenge for the destruction of their city a few decades earlier), or of the Hittites (or a combination of both). If they were S116/P312, it means that they could have been Proto-Celts from Hallstatt. Of course it can't be ruled out that the Trojans asked their "cousins" from Hallstatt for help to defeat the Myceneans, thus invading as a hybrid R1b faction of S116/P312 and ht35. The S116/P312 element could also be due to the later Roman occupation of Greece.
The Cimmerians were the last recorded to leave the Pontic-Anatolian homeland around 800 BCE, passing through Anatolia before going to Europe. The Athenians of Classical Greece (510-323 BCE) made a point to re-established the connections with all the Black Sea ports afterwards, as if to confirm their new genealogical tie with the old Dorian/Trojan homeland (or simply because they could, for the first time in history, since most of the R1b civilization had emigrated).
The Central Asian branch
An early group of R1b1b people is thought to have migrated from Caspian Sea region to Central Asia, where it evolved into the R1b1b1 (M73) branch. This variety of R1b occurs almost exclusively in very specific Central Asian populations. The highest percentages were observed among the Uyghurs (20%) of Xinjiang in north-west China, the Hazara people of Afghanistan (32%), and the Bashkirs (55%) of the Abzelilovsky district of Bashkortostan in Russia (border of Kazakhstan).
Central Asian R1b1b1 could correspond to the Tocharian branch of the Indo-Europeans. It is possible that the Tocharians split from the main R1b body as early as 7,000 BCE. Over the centuries some groups of these nomadic tribes ended up around the southern Urals, others in the Tarim Basin (Xinjiang) or in southern Central Asia.
Mummies of fair-haired Caucasian people were found in the Tarim Basin, the oldest of which date back to 1800 BCE. The modern inhabitants of the Tarim Basin, the Uyghurs, belong both to this R1b-M73 subclade (about 20%) and to R1a1 (about 30%). This could mean that they had become a hybrid R1b-R1a society by the time they reached the Tarim Basin. But R1a1 could also have arrived independently during the Indo-Iranian migrations, or much later through some nomadic Scytho-Iranian tribes.
The earliest known back migration of R1b was from Asia to Africa and took place around 15,000 years ago. A group of R1b1* people moving from the Levant to Egypt, Sudan and spreading in different directions inside Africa to Rwanda, South Africa, Namibia, Angola, Congo, Gabon, Equatorial Guinea, Cameroon, Nigeria, Ivory Coast, Guinea-Bissau. The hotspot is Cameroon. R1b1* was observed at a frequency of up to 95% in some tribes of northern Cameroon (like the Kirdi), and about 15% nationwide. It is in all likelihood where the early R1b people first settled, then spread south and east along the coast.
Other back migrations occured from Europe to the Near East and Central Asia during the Antiquity and Middle Ages. R1b-S28 was found in Romania, Turkey and at the border of Kazakhstan and Kyrgyzstan. Some of it was surely brought by the Alpine Celts (Hallstatt/La Tène culture), known to have advanced along the Danube, and created the Galatian kingdom in central Anatolia. The rest could just as well be Roman, given that R1b-S28 is the dominant form of R1b in the Italian peninsula. Some have hypothetised that Roman legions went as far as Central Asia or China and never came back, leaving their genetic marker in isolated pockets. See also Were the Romans and the Alpine Celts close cousins ?
A small percentage of Western European R1b subclades were also found among Christian communities in Lebanon. They are most likely descendants of the crusaders.
How did R1b come to replace most of the older lineages in Western Europe ?
Until recently it was believed that R1b originated in Western Europe due to its strong presence in the region today. The theory was that R1b represented the Paleolithic Europeans (Cro-Magnon) that had sought refuge in the Franco-Cantabrian region at the peak of the last Ice Age, then recolonised Central and Northern Europe once the ice sheet receded. The phylogeny of R1b proved that this scenario was not possible, because older R1b clades were consistently found in Central Asia and the Middle East, and the youngest in Western and Northern Europe. There was a clear gradient from East to West tracing the migration of R1b people (see map above). This age of the main migration from the shores of the Black Sea to Central Europe also happened to match the timeframe of the Indo-European invasion of Europe, which coincides with the introduction of the Bronze-Age culture in Western Europe, and the spread of Italo-Celtic and Germanic languages.
Historians and archeologists have long argued whether the Indo-European migration was a massive invasion, or rather a cultural diffusion of language and technology spread only by a small number of incomers. The answer could well be "neither". Proponents of the diffusion theory would have us think that R1b is native to Western Europe, and R1a alone represent the Indo-Europeans. The problem is that haplogroup R did arise in Central Asia, and R2 is still restricted to Central and South Asia, while R1a and the older subclades of R1b are also found in Central Asia. The age of R1b subclades in Europe coincide with the Bronze-Age. R1b must consequently have replaced most of the native Y-DNA lineages in Europe from the Bronze-Age onwards.
However, a massive migration and nearly complete anihilation of the Paleolithic population can hardly be envisaged. Western Europeans do look quite different in Ireland, Holland, Aquitaine or Portugal, despite being all regions where R1b is dominant. Autosomal DNA studies have confirmed that the Western European population is far from homogeneous. A lot of maternal lineages (mtDNA) also appear to be of Paleolithic origin (e.g. H1, H3, U5 or V) based on ancient DNA tests. What a lot of people forget is that there is also no need of a large-scale exodus for patrilineal lineages to be replaced fairly quickly. Here is why.
Polygamy. Unlike women, men are not limited in the number of children they can procreate. Men with power typically have more children. This was all the truer in primitive societies, where polygamy was often the norm for chieftains and kings.
Status & Power. Equipped with Bronze weapons and horses, the Indo-Europeans would have easily subjugated the Neolithic farmers and with even greater ease Europe's last hunter-gatherers.If they did not exterminate the indigenous men, the newcomers would have become the new ruling class, with a multitude of local kings, chieftains and noblemen (Bronze-Age Celts and Germans lived in small village communities with a chief, each part of a small tribe headed by a king) with higher reproductive opportunities than average.
Gender imbalance. Invading armies normally have far more men than women. Men must therefore find women in the conquered population. Wars are waged by men, and the losers suffer heavier casualties, leaving more women available to the winners.
Aggressive warfare. The Indo-Europeans were a warlike people with a strong heroic code emphasising courage and military prowess. Their superior technology (metal weapons, wheeled vehicles and warhorses) and attitude to life would have allowed them to slaughter any population that did not have organised armies with metal weapons (i.e. anybody except the Middle-Eastern civilizations).
Replacement of patrilineal lineages following this model quickly becomes exponential. Imagine 100 Indo-European men conquering a tribe of 1000 indigenous Europeans (a ratio of 1:10). War casualties have resulted in a higher proportion of women in the conquered population. Let's say that the surviving population is composed of 700 women and 300 men. Let's suppose that the victorious Indo-European men end up having twice as many children reaching adulthood as the men of the vanquished tribe. There is a number of reason for that. The winners would take more wives, or take concubines, or even rape women of the vanquished tribe. Their higher status would garantee them greater wealth and therefore better nutrition for their offspring, increasing the chances of reaching adulthood and procreating themselves. An offspring ratio of 2 to 1 for men is actually a conservative estimate, as it is totally conceivable that Bronze-Age sensibilities would have resulted in killing most of the men on the losing side, and raping their women (as attested by the Old Testament). Even so, it would only take a few generations for the winning Y-DNA lineages to become the majority. For instance, if the first generation of Indo-Europeans had two surviving sons per man, against only one per indigenous man, the number of Indo-European paternal lineages would pass to 200 individuals at the second generation, 400 at the third, 800 at the fourth and 1600 at the fifth, and so on. During that time indigenous lineages would only stagnate at 300 individuals for each generation.
Based on such a scenario, the R1b lineages would have quickly overwhelmed the local lineages. Even if the Indo-European conquerors had only slightly more children than the local men, R1b lineages would become dominant within a few centuries. Celtic culture lasted for over 1000 years in Continental Europe before the Roman conquest putting an end to the priviledges of the chieftains and nobility. This is more than enough time for R1b lineages to reach 50 to 80% of the population.
The present-day R1b frequency forms a gradient from the Atlantic fringe of Europe (highest percentage) to Central and Eastern Europe (lowest), the rises again in the Anatolian homeland. This is almost certainly because agriculture was better established in Eastern, then Central Europe, with higher densities of population, leaving R1b invadors more outnumbered than in the West. Besides, other Indo-Europeans of the Corded Ware culture (R1a) had already advanced from modern Russia and Ukraine as far west as Germany and Scandinavia. It would be difficult for R1b people to rival with their R1a cousins who shared similar technology and culture. The Pre-Celto-Germanic R1b would therefore have been forced to settled further west, first around the Alps, then overtaking the then sparsely populated Western Europe.
Distribution of haplogroup R1b in Europe
Subclades of R1b
Defining mutation
Subclade (previous name)
Time of origin (approximate)
Place of highest frequency
Most prevalent ancient ethnic group
M18
R1b1a (R1b1a)
11,000 ybp
Levant, Sardinia
Phoenician, Druze
M73
R1b1b1 (R1b1b)
9,500 ybp
Central Asia
Tocharian
M269
R1b1b2 (R1b1c)
9,500 ybp
Western Europe
Italo-Celto-Anatolian
L23/S141
R1b1b2a
7,000 ybp
Western Europe
Italo-Celto-Anatolian
L11/S127, P311/S128, P310/S129
R1b1b2a1
5,000 ybp
Western Europe
Italo-Celto-Germanic
M405/S21/U106
R1b1b2a1a
(R1b1c9)
3,000 ybp
Frisia, Benelux, England, Austria, northern Italy
West Germanic (Frisian, Anglo-Saxon, Lombard)
M467/S29/U198
R1b1b2a1a1
(R1b1c9b)
1,800 ybp
Southern England + northern Germany
Germanic (Anglo-Saxon)
P107
R1b1b2a1a2
1,800 ybp
Germanic
L1/S26
R1b1b2a1a3
(R1b1c9a)
1,800 ybp
Southern & eastern England, Norway, southern Germany,
and Spain
Germanic
L48
R1b1b2a1a4
Germanic
L44 => L47
R1b1b2a1a4a
Germanic
L5
R1b1b2a1a5
1,800 ybp
Germanic
L6
R1b1b2a1a6
1,800 ybp
Germanic
P312/S116
R1b1b2a1b
4,500 ybp
Western Europe
Italo-Celtic
M65
R1b1b2a1b1
(R1b1c2)
3,500 ybp
Basque country
Basque
M153
R1b1b2a1b2
(R1b1c4)
3,350 ybp
Basque country and Gascony
Basque
M167/SRY2627
R1b1b2a1b3
(R1b1c6)
2,850 ybp
Spain (esp. Catalonia), Western France, Cornwall, Wales
Basque, Catalan, Gascon, Breton, Cornish
S28/U152
R1b1b2a1b4
(R1b1c10)
3,500 ybp
Rhine & Meuse region, Alps, northern Italy
Alpine Celts (Hallstatt-La Tène), Italics
M126
R1b1b2a1b4a
(R1b1c3)
2,500 ybp
Found in Germany, England and Ireland
Celtic
M160
R1b1b2a1b4b
(R1b1c5)
2,000 ybp
Found in Germany and Switzerland
Alpine Celtic
L2/S139
R1b1b2a1b4c
2,500 ybp
Found in Italy, Germany, Belgium, Britain, Ireland, Norway
Alpine Celtic
L20/S144
R1b1b2a1b4c1
1,800 ybp
Found in England, France and Italy
Alpine Celtic
M228
R1b1b2a1b4c1a
Found in northern Italy
Alpine Celtic
L3
R1b1b2a1b4c2
Found in Germany and England
Alpine Celtic
L4
R1b1b2a1b4d
Found in Latvia and Poland
S68
R1b1b2a1b5
(R1b1c11)
3,500 ybp
Sweden and Scotland
Germanic
L21/S145
R1b1b2a1b6
4,000 ybp
Ireland, Britain, France, Germany
Celtic, Brythonic, Gaelic
M37
R1b1b2a1b6a
(R1b1c1)
3,000 ybp
Ireland, Britain, France, Germany
Celtic
M222
R1b1b2a1b6b
(R1b1c7)
3,000 ybp
North-west Ireland and west Scotland
Scottish Irish
P66
R1b1b2a1b6c
(R1b1c8)
2,500 ybp
?
?
L7
R1b1b2a1b7
?
?
L8
R1b1b2a1b8
?
?
L9, L10
R1b1b2a1b9
?
?
Haplogroup R1a (Y-DNA)
R1a is thought to have been the dominant haplogroup among the Indo-European speakers who evolved into the Indo-Iranian, Mycenaean Greek, Macedonian, Thracian, Baltic and Slavic branches.
The most likely origin of the Proto-Indo-Europeans lies in the Kurgan culture (7000-3000 BCE) and the subsequent Yamna culture (3500-2200 BCE), between modern Ukraine and south-west Russia. Their expansion is linked to the domestication of horses in the Eurasian steppes, and the invention of the chariot (see R1b above).
The western branch
The first expansion of R1a took place with the westward diffusion of the Corded Ware (or Battle Axe) culture (3200-1800 BCE) from the Yamna homeland. This resulted in R1a being the dominant haplogroup in the northern half of Eastern Europe nowadays, like in Poland (56% of the population), Ukraine (50%), Belarus (45%), Russia (40%), Slovakia (40%), Latvia (40%), Lithuania (38%) and the Czech Republic (34%)
The Germanic branch of Indo-European languages probably inherited more from the R1b cultures, although R1a is likely to have arrived earlier in Scandinavia, during the Corded Ware period. R1a people would have mixed with the pre-Germanic I1 aborigines to create the Nordic Bronze Age (1800-500 BCE). R1b would have reached Scandinavia later as a northward migration from the contemporary Hallstatt culture (1200-500 BCE). The first truly Germanic tongue could have been a blend of Hallstatt Proto-Celtic and the Corded-Ware Proto-Slavic with a few pre-Germanic loan words. The fact that present-day Scandinavia is composed of roughly 40% of I1, 20% of R1a and 40% of R1b reinforces the idea that Germanic ethnicity and language had acquired a tri-hybrid character by the Iron Age.
The Corded Ware culture was followed by the Trzciniec culture (1700-1200 BCE) and the Lusatian culture (1300-500 BCE) in modern Balto-Slavic countries.
Historically, no other part of Europe was invaded a higher number of times by R1a steppe peoples than the Balkans. Chronologically, the first R1a invaders came with the westward expansion of the Corded Ware culture (from about 3200 BCE), then the Mycenaean invasion (1600 BCE), followed by the Thracians (1500 BCE), the Illyrians (around 1200 BCE), the Huns and the Alans (400 CE), the Avars, the Bulgars and the Serbs (all around 600 CE), and the Magyars (900 CE), among others. These peoples originated from different parts of the Eurasian steppes, anywhere between Eastern Europe and Central Asia, which is why such high STR diversity is found within Balkanic R1a nowadays. It is not yet possible to determine the ethnic origin for each variety of R1a, apart from the fact that about any R1a is associated with tribes from Eurasian steppe at one point in history.
The eastern branch
The eastern branch of the R1a steppe people was the Andronovo culture (2300-1000 BCE), around modern Kazakhstan, which correspond to the Indo-Iranian branch of languages. Their migration to the south have resulted in high R1a frequencies in southern Central Asia, Iran and the Indian subcontinent. The highest frequency of R1a (about 65%) is reached in a cluster around Kyrgyzstan, Tajikistan and northern Afghanistan. In India, 15 to 45% of the population is R1a, depending on the region and caste. Over 70% of the Brahmins (highest caste in Hindusim) belong to R1a1, due to a founder effect.
Distribution of haplogroup R1a in Eurasia
Haplogroup I (Y-DNA)
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 around 35,000 years ago, and developed into haplogroup I approximately 25,000 years ago. This means that Cro-Magnons most probably belonged (exclusively ?) to IJ or I. Nowadays haplogroup I accounts for 10 to 45% of the population in most of Europe. It is divided in four main subclades.
The megalithic structures (5000-1200 BCE) of Europe were built by I people.
Haplogroup I1 (formerly I1a) is the most common I subclade. It is found mostly in Scandinavia and Northern Germany, where it can represent over 35% of the population. Associated with the Norse ethnicity, it is found in all places invaded by the ancient Germanic tribes and the Vikings.
During the Neolithic period, pre-I1 and I1 people were part of the sucessive Ertebølle culture (5300-3950 BCE) and Funnelbeaker culture (4000-2700 BCE). The Corded Ware period (3200-1800 BCE) marks the arrival of the Indo-European R1a people from the Ukrainian steppes.
I1 is identified by at least 15 unique mutations, which indicates that this lineage has been isolated for a long period of time, or experienced a serious population bottleneck. Although the first mutation splitting I1 away from I2 may have arisen as long as 20,000 years ago, people belonging to this haplogroup all descend from a single man who lived less than 5,000 years ago. This corresponds to the arrival of the Indo-European, suggesting that a high percentage of the indigenous I1 men could possibly have been killed by the new immigrants.
Distribution of haplogroup I1 in Europe
Haplogroup I2 might have originated in southeastern Europe some 17,000 years ago and developed into four main subgroups : I2a1, I2a2, I2b1 and I2b2.
I2a1 (formerly I1b2) is found chiefly among the Sardinians and the Basques, and is rarely found outside Iberia, Western France, the West coast of Italy and the Mediterranean coast of the Maghreb. It accounts for approximately 40% of all Y-DNA haplogroups among the Sardinians. I2a1 is estimated to be 8,000 years old.
I2a2 (formerly I1b) is typical of the Dinaric Slavs (Croats, Serbs and Bosniaks). Its highest density is observed around ex-Yugoslavia and Moldova, but it is also common to a lower extent in Albania, Northern Greece, Bulgaria, Romania, Ukraine, Belarus, and southwestern Russia.
The Neolithic period in I2a1 and I2a2 territory (Illyria, Italy, Sardinia, Mediterranean coast of France and Spain) matches the Printed-Cardium Pottery culture (5000-1500 BCE), that is believed to have started with the arrival of E-V13 and G2a farmers and herders from Thessaly (northern Greece). It was followed by the Terramare culture (1500-1000 BCE) in the Bronze Age. The R1b Celto-Italic people are thought to have crossed the Alps and invaded the Italian peninsula around 1,000 BCE, replacing most of the indigenous I2a, G2a and E-V13 people (especially in the northern half).
I2b (formerly I1c) is associated with the pre Celto-Germanic people of North-Western Europe, such as the megaliths builders (5000-1200 BCE). The wide variety of STR markers within I2b could make it as much as 13,000 years old.
I2b is found in all Western Europe, but apparently survived better the Indo-European invasions (=> see R1b above) in northern Germany, and was reintroduced by the Germanic invasions during the late Roman period. Nowadays, I2b peaks in central and northern Germany (10-20%), the Benelux (10-15%) as well as in northern Sweden. It is also found in 3 to 10% of the inhabitants of Denmark, East England, and Northern France. It is rare in Norway, which concords with the fact that it hasn't been invaded by people from northern Germany.
There are two major subclades : I2b1 (M223+) and I2b2 (L38/S154+), further subdivided in at least 4 subclades each, although little is known about them yet. The subclade I2b1a (M284+) occurs almost exclusively in Britain, where it seemingly developed about 3,000 years ago.
Near-Eastern haplogroups
Haplogroup J (Y-DNA)
J is a Middle Eastern haplogroup, divided into the northern J2 and the southern J1. J2 is by far the most common variety in Europe.
J2 originated in northern Mesopotamia, and spread westward to Anatolia and southern Europe, and eastward to Persia and India. J2 is related to the Ancient Etruscans, (Minoan) Greeks, southern Anatolians, Phoenicians, Assyrians and Babylonians.
In Europe, J2 reaches its highest frequency in Greece (especially in Crete, Peloponese and Thrace), southern and central Italy, southern France, and southern Spain. The ancient Greeks and Phoenicians were the main driving forces behind the spread J2 around the western and southern Mediterranean.
J2 is thought to have arrived in Greece from Anatolia in the early Neolithic, or possibly even earlier. J2b perhaps originated in Greece (or in Anatolia ?), like haplogroup E-V13 (see below) to which it is closely linked. The propagation of J2b and E-V13 (as well as a minority of T) follows the diffusion of agriculture across the Balkans, the Danube basin, and until the north of France to the west, and Moldova to the east. Apart from south-east Europe, J2b is also found all around India, but only at moderate levels in between Europe and India.
The world's maximum concentrations of J2a is in Crete (32% of the population). The subclade J2a8 appears to be native to Crete. J2a also reaches high frequencies in Anatolia and the southern Caucasus. A likely place of origin is northern Mesopotamia.
Interestingly, J2a* is found as far as India and is largely confined to the upper castes. The Brahmin (priest) caste is made up almost exclusively of haplogroups R1a1, R2, and J2a (although R1a1 makes up two thirds of the lineages). These 3 haplogroups have Bronze Age coalescence time and are thought to represent the gene flow of the Indo-Aryan invasion of the Indian subcontinent about 3,500 years ago.
J1 is a typically Semitic haplogroup, making up most of the population of the Arabian peninsula. Its highest density is observed in Yemen (72%), which could be its native place. The Muslim conquest of the Middle East, North Africa, and to a lower extent also to Sicily and southern Spain, spread J1 far beyond Arabia, creating a new Arabic world.
A considerable part of Jewish people belong to J1 and J2, although J2 is more common. J1 is the Cohen Modal Haplotype, meaning that about three quarters of the people called Cohen, Kohen, or a variant belong to a specific J1 haplotype. In the Hebrew Bible the common ancestor of all Cohens is identified as Aaron, the brother of Moses.
Haplogroup E1b1b (Y-DNA)
Haplogroup E1b1b (formerly E3b) represents the last major migration out of Africa into Europe. It is believed to have first appeared in the Horn of Africa or southern Africa approximately 26,000 years ago and dispersed to the Middle East during the Upper Paleolithic and Mesolithic periods.
On the European continent it has the highest concentration in north-west Greece, Albania and Kosovo, then fading around the Balkans, the rest of Greece and Western Turkey. Outside Europe, it is also found in most of the Middle East, northern and eastern Africa, especially in Morocco, Lybia, Egypt Yemen, Somalia, Ethiopia and South Africa.
The vast majority of Europeans and Near Easterners belong to E1b1b1a (or E-M78, formerly E3b1a). E-M78 is thought to migrated out of Egypt in the early Neolithic to colonise the Levant, Anatolia and Greece, where it mixed with the J2 inhabitants. E-M78 then evolved into 4 main branches : E1b1b1a1 (E-V12), E1b1b1a2 (E-V13), E1b1b1a3 (E-V22) and E1b1b1a4 (E-V65), each further subdivided in "a" and "b" subclades.
E-M78 from the Near East settled in northern Greece circa 8,500 years ago, launching the Thessalian Neolithic (6500-2500 BCE). E-V13 could have originated in Thessaly around 8,000 years ago, before expanding towards the Balkans. In the following millennia, the expansion of E-V13 correlates with the spread of Linear Pottery culture (5500-4500 BCE) from the Balkans to Central Europe and Germany. Owing to this early introduction to Europe, E-V13 is now by far the most common E subclade found among Europeans.
E-V13 is also associated with the ancient Greek expansion and colonisation. Outside of the Balkans and Central Europe, it is particularly common in Southern Italy, Cyprus and Southern France, all part of the Classcial ancient Greek world.
E-V22 is the predominant subclade in the Levant and is therefore associated with the Phoenicians and Jews. It is also common in Egypt, where it might have originated. The Phoenicians spread E-V22 to Sicily, Sardinia, southern Spain and the Maghreb, and the Jews to Spain and Italy. Approximately half of Spanish and Italian E are V-22 (Jewish-Phoenician), and the other half V-13 (Greek).
E-V12 is the most common subclade of M78 in Egypt. Its low presence around Greece and Anatolia indicates that it probably already existed when E moved there in the early Neolithic.
E-V65 is found in North Africa, with a maximum frequency in Lybia, then Morocco. It is also likely to have originated in Egypt. In Europe it is found at low frequencies in Greece and Sicily, but interestingly makes up one fourth of Sardinian E. It could be due to immigration from the Phoenician colonies in the Maghreb to Sardinia (the Sardinian haplogroup I2a1 is also present at low frequencies along the coast of Algeria and Tunisia, confirming exchanges of population between the two regions, maybe when both were Phoenician colonies).
E1b1b1b (E-M81, formerly E3b1b) is characteristic of the Berbers of North-West Africa. In some parts of Morocco E1b1b1b can peak at 80% of the population. This sub-hapolgroup is also found in Iberia, Italy and southern France, with the highest concentrations in southern Portugal (12%) and decreasing as we move north.
Expansion of haplogroup E from Africa to Europe from the pre-Neolithic to the Phoenician colonization (9500-800 BCE)
Haplogroup G (Y-DNA)
G has its roots in around the Caucasus. It is found mostly in mountainous regions between the Near East and India (Caucasus, Iran, Afghanistan, Kashmir), but also in Central Asia (Kazakhstan), Europe and North Africa.
Most Europeans belong to the G2a subclade, and most northern and western Europeans more specifically to G2a3b (or to a lower extend G2a3a). About all G2c Europeans are Ashkenazi Jews. The discovery of G2c subclades around Afghanistan indicates that it could have originated in that part of the world. G1 is found predominantly in Iran, but is also found in Central Asia (Kazakhstan). A famous members of haplogroup G was Joseph Stalin (G2a1), who was of Georgian origin.
There are several theories regarding the origin of G2a in Europe. There are doubtlessly cumulative rather than exclusive.
Neolithic hypothesis
G2a makes up 5 to 10% of the population of Mediterranean Europe, but is fairly rare in Northern Europe. The only places where haplogroup G2 exceeds 10% of the population in Europe are Switzerland, the Tyrol, south-central Italy (Molise, Central and Southern Apennine), Sardinia, northern Greece (Thessaly) and Crete - all mountainous and relatively isolated regions.
Chronologically, the first hypothesis is the advance of Neolithic farmers and herders from Anatolia to Europe between 9,000 and 6,000 years ago. It is likely that these Caucasian migrants brought with them sheep and goats, which were domesticated south of the Caucasus arbout 12,000 years ago. This may explain why haplogroup G is more common in mountainous areas, be it in Europe or in Asia.
Haplogroup G has also been linked to the spread of metalworking from the Caucasus or Anatolia to places like Sardinia. The age of G2a combined with geographic continuity of G2a from Anatolia to Thessaly to the Italian peninsula, Sardinia, south-central France and eastern Spain strongly suggest that G2a is connected to the Printed-Cardium Pottery culture (5000-1500 BCE).
A Neolithic introduction of G to Europe would undeniably correspond to G2a3, which is by far the most common and diversified subclade in Europe.
Roman hypothesis
It is most likely that G2a arrived in Europe during the Neolithic and that the Romans helped spread it around. Migrations within the Roman Empire probably contributed to a moderate increase of G2a northward to Gaul and Britain, Indeed, the frequency of haplogroup G decreases with the distance from the boundaries of the Roman Empire, Haplogroup G is extremely rare Nordic and Baltic countries nowadays, despite the fact that agriculture reached those regions around the same time as Britain or Ireland.
Alanic hypothesis
The only ethnic group that has a majority of haplogroup G nowadays are the Ossetians in the Caucasus, in the modern Russian Republic of North Ossetia-Alania. They are thought to descend directly from the Alans, a Central Asian tribe related to the ancient Samartians. The medieval Kingdom of Alania was located in the northern Caucasus, in present-day Georgia and Ossetia.
G2a has been observed at a slightly higher frequencies in Picardy and Flanders than in surrounding regions. It has been hypothetised that G2a was brought to northern France and Belgium by the Alans, who traversed all continental Europe during the barbarian invasions in the 5th century and founded a short-lived kingdom in northern France.
Nonetheless, if there is Alanic G in Europe it must certainly belong to other subclades than those from the Neolithic period (namely G2a3). G2a1 being the most common variety in the Caucasus nowadays, the fairly recent Alanic migration (from a genetic point of view) could have carried that particular subclade. In fact, G2a1 has been found all along the Alanic migration route (Hungary, France, Spain), as well as in Britain (Samartian element ?), but hardly anywhere else.
Scythian hypothesis
Romans were known to recruit Scythian or Sarmatian horsemen in their legions. According to C. Scott Littleton in his book From Scythia to Camelot, several Knights of the Round Table were of Scythian origin, and the the legend of Holy Grail itself originated in ancient Scythia. This hypothesis was also taken up in the 2004 movie King Arthur, which opens with the arrival of Scytho-Roman cavalry in Britain. However, Scythians were steppe people more likely to belong to haplogroup R1a. If any of them did belong to G, they presumably were G1, not G2a. This would explain the few cases of G1 in north-western Europe though.
Haplogroup T (Y-DNA)
T is a rare haplogroup in Europe (less than 1% of the population). It originated around the Red Sea (maybe in Ethiopia) at least 30,000 years ago, making it one of the oldest haplogroups found in Eurasia. It is most common in north-east Africa and the west coast of the Arabian peninsula, where it accounts for approximately 5 to 8% of the male lineages. Besides these regions and Europe, T is found as far as southern India, Russia, Tanzania and Cameroon. Its highest density is actually found among the Fulbe people of Cameroon (18% of the population).
Within Europe there are a few pockets with surprisingly high densities of haplogroup T, like the town of Sciacca in Sicily (18%), on the Spanish island of Ibiza (17%) or Serbia (7%). The populations of Italy, Portugal, Greece, and (oddly enough) Estonia, all have between 3 and 4% of haplogroup T.
The spread of haplogroup T in Europe is closely linked to the expansion of E1b1b from Egypt and the Near East to the Balkans and Danube basin. Its presence around the Mediterranean can be attributed to the Phoenicians colonisation (1200-800 BCE). The pocket in Estonia might be due to a founder effect in the region's Jewish population. Among famous people, Thomas Jefferson belonged to haplogroup T.
Other haplogroups found in Europe
Haplogroup N (Y-DNA)
N is found among Uralic speakers, from Finland to Siberia, and at minor frequencies as far as Korea and Japan. In Europe, haplogroup N is only found at high frequencies among modern Finns (58%), Lithuanians (42%), Latvians (38%), Estonians (34%) and northern Russians.
Haplogroup N is believed to have originated in Southeast Asia approximately 15,000 to 20,000 years ago, but the N1c1 subclade found in Europe likely arose in Southern Siberia circa 12,000 years ago, and spread to North-East Europe 10,000 years ago.
Haplogroup N is associated with the Kunda culture (8000-5000 BCE) and the Comb Ceramic culture (4200-2000 BCE), which evolved into Finnic and pre-Baltic people.The Indo-European Corded Ware culture (3200-1800 BCE) progressively took over the Baltic region and southern Finland from 2,500 BCE. The merger of the two gave rise to the hybrid Kiukainen culture (2300-1500 BCE). Modern Baltic people have a roughly equal proportion of haplogroup N1c1 and R1a, resulting from this merger of Uralic and Slavic cultures.
Haplogroup Q (Y-DNA)
Q is thought to be the dominant haplogroup of the Huns, who invaded Europe in the 5th century, and is only found in 2% of the people in Hungary, where the one Hunnic tribe finally settled. Another group of Huns could have settled in Sweden and/or Norway, where Q is also found in among 0.5% of the population.
Haplogroup C (Y-DNA)
Haplogroup C3 in Europe is most likely of Mongol origin. It is found everywhere at various concentrations in Genghis Khan's former empire, although only sporadically on the European continent. Other subclades of C come from ethnic groups too remote from Europe (Aboriginal Australians, Polynesians, South-East Asians) to be found among Europeans (apart from recent immigration).
Haplogroup P (Y-DNA)
P is the parent group of Q and R (including R1a and R1b). It has almost disappeared nowadays, except around its place of origins in Central Asia. It is very rarely found in Europe. It may have been brought to Europe by Central Asian invaders, like the Huns or the Mongols.
Haplogroup L (Y-DNA)
L is found mostly in the Indian subcontinent, but also at lower frequencies in Central Asia, Southwest Asia, and Southern Europe along the coast of the Mediterranean Sea (notably in Italy). L1 is typical of the Dravidian people of South India. Various subclades are found in Europe (L1, L2, L3) without any real geographic pattern. Europeans belonging to haplogroup L are likely to be descended from Indian (L1, L3) or Persian (L2, L3) merchants in ancient times, maybe at the time of the Roman Empire.
Haplogroup H (Y-DNA)
Gypsies belong predominantly (about 50%) to haplogroup H1a. Haplogroup H is not otherwise found in Europe, but on the Indian subcontinent.
Haplogroup A (Y-DNA)
A is the oldest of all Y-DNA haplogroups and the closest to the Y-chromosomal Adam. It originated in Africa over 70,000 years ago, most likely in the south-west, around modern Angola and Namibia. Modern populations with the highest percentages of haplogroup A are the Khoisan (such as the Bushmen) and the southern Sudanese. Isolated cases of individuals belonging to haplogroup A have been found in Western Europe (notably Ireland, Britain and Germany). It is believed that these people descend in direct paternal line from Nubians who probably came to Europe during the Roman period, probably as slaves (Nubian gladiators were popular in Rome). It is unlikely that they descend from slaves from the Atlantic slave trade (17th and 18th century) since these came from a part of Africa where A is very rare.
MtDNA Haplogroups
All mtDNA haplogroups found in Europe descend from the N group, which is thought to represent one of the two initial migrations by modern humans out of Africa, some 60,000 to 80,000 years ago. Nowadays haplogroup N is only found at extremely low frequencies in various parts of Eurasia.
Unfortunately, the tiny size of mitochondrial DNA (approximately 16,500 base pairs as opposed to 60 million for Y-DNA) does not allow a very accurate tracing of ancestry. Mitochondrial haplogroups all arose during the Ice Age, a period when humans were nomadic hunter-gatherers, well before the establishment of cities and civilizations. Mitochondrial haplogroups are only linked to ethnicities at a continental level. Those associated with European descent are H, I, J, K, T, U, V, W and X (except the branch X2a which found among Native Americans). Deep subclades can be associated with more specific regions, but do not necessarily match historical ethnic and linguistic groups. One likely reason is that women, through whom mtDNA is passed, tended to marry outside their ethnic group more often than men (e.g. to secure an alliance between two tribes or kingdoms).
Chronological development of mtDNA haplogroups
Note that the age of mitochondrial haplogroups is much more difficult to estimate than Y-DNA haplogroups, due to the tiny sequence of mtDNA and the few number of mutations available. The error margin for the dates below is typically of +-5,000 years, but could even exceed that for older haplogroups.
N => 75,000 years ago (arose in North-East Africa)
R => 70,000 years ago (in South-West Asia)
U => 60,000 years ago (in North-East Africa or South-West Asia)
pre-JT => 55,000 years ago (in the Middle East)
JT => 50,000 years ago (in the Middle East)
U5 => 50,000 years ago (in Western Asia)
U6 => 50,000 years ago (in North Africa)
U8 => 50,000 years ago (in Western Asia)
pre-HV => 50,000 years ago (in the Near East)
J => 45,000 years ago (in the Near East or Caucasus)
HV => 40,000 years ago (in the Near East)
X => over 30,000 years ago (in north-east Europe)
U5a1 => 30,000 years ago (in Europe)
I => 30,000 years ago (origin unknown - probably in Europe)
J1a => 27,000 years ago (in the Near East)
W => 25,000 years ago (in north-east Europe or north-west Asia)
U4 => 25,000 years ago (in Central Asia)
J1b => 23,000 years ago (in the Near East)
H => over 20,000 years ago (in the Near East or Southern Europe)
T => 17,000 years ago (in Mesopotamia)
K => 16,000 years ago (in the Near East)
V => 15,000 years ago (arose in Iberia and moved to Scandinavia)
H1b => 13,000 years ago (in Europe)
K1 => 12,000 years ago (in the Near East)
H3 => 10,000 years ago (in Western Europe)
Mitochondrial DNA of prehistoric Europeans
The testing of ancient DNA helped understand how long each haplogroup has been in Europe. Only a few such tests have been successfully conducted so far. Mitochondrial DNA was extracted from the skeleton of a 28,000 year-old Cro-Magnon from southern Italy, and the haplogroup was determined as HV or pre-HV. Still preceding the Neolithic expansion from the Middle East, the 9,000 year-old Cheddar Man was found to belong to haplogroup U5a. (=> More examples of ancient mtDNA haplogroups).
Autochtonous (Cro-Magnoid) Europeans must have therefore belonged at least to haplogroups HV (and its offspring H and V) as well as U5a, which also happen to be the most common mitochndrial haplogroup everywhere in Europe. It has been speculated that over half of the matrilineal lineages in Europe descend directly from Paleolithic Europeans. Their male counterpart are Y-DNA haplogroup I.
European mtDNA haplogroups and their subclades
Haplogroup H & V (mtDNA)
Haplogroup H is by far the most common all over Europe, amounting to about 40% of the European population. It is also found (though in lower frequencies) in North Africa, the Middle East, Central Asia, Northern Asia, as well as along the East coast of Africa as far as Madagascar.
H1, H3 and V are the most common subclades of HV in Western Europe. H1 peaks in Norway (30% of the population) and Iberia (18 to 25%), and is also high among the Sardinians, Finns and Estonians (16%), as well as Western and Central European in general (10 to 12%) and North-West Africans (10 to 20%). H3 is commonest in Portugal (12%), Sardinia (11%), Galicia (10%), the Basque country (10%), Ireland (6%), Norway (6%), Hungary (6%) and southwestern France (5%). Haplogroup V reaches its highest frequency in northern Scandinavia (40% of the Sami), northern Spain, the Netherlands (8%), Sardinia, the Croatian islands and the Maghreb. It is likely that H1, H3 and V, along with haplogroup U5, were the main haplogroups of Western European hunter-gatherers living in the Franco-Cantabrian refuge during the last Ice Age, and repopulated much of Central and Northern Europe from 15,000 years ago.
Haplogroup H13 is most common in Sardinia and around the Caucasus. Its distribution is reminiscent of Y-DNA haplogroup G2a. The same is true of H2 to a lower extent. This would suggest a Caucasian or Anatolian origin.
H5 and H7 are also common in the Caucasus, but their lower incidence around the Mediterranean, and higher frequency from Anatolia to the Alps via the Danube suggest a possible link with the spread of R1b.
Haplogroup U & K (mtDNA)
Haplogroup U is extremely old. It originated some 60,000 years ago at the confine of North-East Africa and the Middle East, soon after the first Homo Sapiens ventured out of Africa. This is why each of its top-level subclade (U1, U2, U3...) can be seen as a haplogroup in its own right. The main European subclades are U3, U4, U5 and U8/K. U1 is mostly found in the Middle East, U2 in South Asia, U6 in North Africa, U7 from the Near East to India, and the rare U9 from Ethiopia and the Arabian peninsula to Pakistan.
Haplogroup U3 is centered around the Black Sea, with a particularly strong concentration in the north-eastern part. It could be related to the ancient Indo-Europeans, and probably more to R1b than R1a.
Haplogroup U4 are more common in Eastern Europe, Central Asia, northern South Asia (around Tajikistan for U4, and Pakistan for W), which also suggests an affiliation with the Indo-Europeans (correlated to Y-DNA haplogroup R1a). The same is true of haplogroups I, W, T2 and U2e to a lower extent.
Haplogroup U5 is the most common in Western and Northern Europe. DNA tests on ancient skeletons have shown that U5 was the principal mitochondrial haplogroup of Paleolithic and Mesolithic hunter-gatherers in Northern Europe. Ancient DNA tests conducted in Britain, Germany and Scandinavia indicate that the frequency of U5 has progressively declined over time through the Neolithic, Bronze Age, Iron Age and Middle Ages. Nowadays it remains most common in the far north of Europe, where the Mesolithic population has been least affected by subsequent migrations. For instance, 30 to 50% of the Sami people of northern Scandinavia belong to haplogroup U5b (and about 40% to haplogroup V, which is also pre-Neolithic European origin).
Haplogroup K is in fact the main subclade of U8. It is found throughout Europe and Western Asia, as far away as India. Its highest concentration is in North-West and Central Europe, Anatolia and the southern Arabian peninsula. It is believed to have first arisen somewhere between the Near East (Anatolia ?) and northeastern Italy approximately 16,000 years ago (estimates range from 22,000 years to as little as 10,000 years before present). It has the largest number of subclades of any haplogroup in spite of its fairly recent age. Most K1a4, K1a10, K1b, K1c and K2 subclades are typically European. K1a1b1a and K1a9 are found primarily among Ashkenazi Jews. Compared with Y-DNA haplogroups, mtDNA K seems to correlate best with the spread of R1b, J1 and J2. This reinforces the hypothesis of an Anatolian origin.
Haplogroup J & T (mtDNA)
Haplogroup J appears to correlate the most with the spread of agriculture from the Middle East. However, being one of the oldest haplogroups, some branches could very well have been in Europe long before that. J1 and J2a are generally more common in Europe, while J2b is more frequent in the Middle East and in South-East Europe.
Haplogroup T is thought to have originated in Mesopotamia some 12,000 years ago. It is found from Europe to India, with the highest concentrations around the eastern Baltic Sea. Like haplogroup J, T probably spread early with Neolithic farmers. The Indo-Europeans may also have help spread it to Europe and South Asia.
Haplogroup X (mtDNA)
Haplogroup X is a very old and scattered haplogroup found all over Eurasia, North Africa as well as among Native North Americans. It frequency rarely exceeds 5% of the population in any ethnic group, and is more often restricted to 1 or 2%. X1 is found almost exclusively in North Africa, while X2b is the only lineage present among Amerindians. X2a, X2c, X2d and X2e are found in Europe, Siberia and Central Asia.
The strong presence of X2 around the Caucasus, progressively fading towards the Near East and Mediterranean , hints that it could be related to the spread of Y-DNA haplogroup G2a.
Finno-Uralic mtDNA
Finno-Uralic people have an overall mtDNA admixture similar to other Europeans, with a higher percentage of W and U5b, and a small percentage of Siberian haplogroups such as N or A. The Sami are characterised by a high percentage of haplogroups U5b1 and V.
Berber mtDNA
The Berbers are the indigenous populationof north-west Africa. Although their Y-DNA is almost perfectly homogenous, belonging to haplogroup E-M81, Berber maternal lineages show a much greater diversity, as well as regional disparity. At least half (and up to 90% in some regions) of the Berbers belong to some Eurasian lineages, such as H, HV, R0, J, T, U, K, N1, N2, and X2, mostly of Middle or Near Eastern origin. 5 to 45% of the Berbers will have sub-Saharan mtDNA (L0, L1, L2, L3, L4, L5). There are only three native North African lineages, U6, X1 and M1, representing 0 to 35% of the people depending on the region.
Haplogroup U6 has been observed from the Iberia and the Canary Islands to Senegal in the West, and from Syria to Ethiopia and Kenya in the East. It is also found at low density in Europe, though mostly limited to Iberia. Approximately 10% of all North Africans belong to this lineage.
Gypsy mtDNA
The Gypsies (Romani people) originated in the Indian subcontinent and mixed with local population in the Middle East and Eastern Europe over the centuries. About half of the Gypsy population belong to haplogroup M, and more specifically M5, which is otherwise exclusive to South Asia. The other mtDNA haplogroups found among the Gypsy community are mostly of Eastern European, Middle Eastern or Central Asian origin, and notably include H, U3, J, I, W and X, depending on the Romani group.
European mtDNA haplogroups and their subclades
Subclade
Time of origin
Place of highest frequency
Most prevalent ancient ethnic group
H1
Spain, Scandinavia, Germany and Russia
H1b
13,000 years ago
Eastern Europe and North Central Europe
Slavic-Germanic
H2
Eastern Europe, Russia, Central Asia, Middle East
H2a
Eastern Europe, Caucasus, and Central Asia
Scythian
H2b
? (Cambridge Reference Sequence)
H3
10,000 years ago
Iberia, Sardinia and Germanic countries
H4
Iberia, Central and South-East Europe
H5
France, Northern Italy, Iberia, Central European plain, Finland
H5a
7,500 years ago
Central Europe
H6
40,000 years ago
Ireland, Central Europe, Russia, Caucasus, Middle East Central Asia
H7
Russia, Central Asia, Caucasus
H8
Central Europe, Syria, Armenia, Central Asia
H13
Southern Europe, Middle East, Caucasus, Russia,
H14
Middle East and Caucasus
H18
Arabian peninsula
Arab
H19
Caucasus
H21
Caucasus
V
15,000 years ago
Basque country and northern Scandinavia
Basque, Saami
U1
India, Middle East, eastern and southern Europe
Near-Eastern
U2
South and Central Asia
Indo-Aryan
U3
Balkans, Anatolia, Caucasus, Middle-East
U4
25,000 years ago
Baltic, Russia, Central Asia
Kurgan (Aryan)
U5
55,000 years ago
Northern & Eastern Europe
U5a
Most of Europe
U5a1
30,000 years ago
Finland and Russia
U5b
Germany, Finland and Russia
U5b1
Nordic countries and Russia
Saami
U6
60,000 years ago
North-West Africa, Iberia, Canary Islands
North-West African
U7
Gujarat (India), Iran, Pakistan, Iran, Near-East and Italy
Persian, North-West Indian
U8
Most of Europe and the Middle East
U8a
Basque country
Basque
U8b
Italy (+ Jordan)
K*
16,000 years ago
Europe & Middle East
K1
Northern Europe, Alps, Italy
K1a
4,000 years sgo
Northern Italy, Alps and Rhine region
K1a1b1a
Eastern Europe & Russia
Ashkenazi Jewish
K1a4
Around the Alps, Germany, Britain and Ireland
Celtic
K1a9
Eastern Europe & Russia
Ashkenazi Jewish
K1a10
North-Western Europe
K1b
Most of Europe
K1c
Most of Western and Northern Europe, especially Iberia
K1c2
Germanic countries
Germanic
K2
Around the Alps
K2a
Eastern, Central and North-Western Europe
Slavic, Germanic or Celtic
K2b
Britain, Ireland, Iberia and the Alps
Celtic
J*
45,000 years ago
Ireland, Britain and Germany
J1
South-East Europe and the Alps
J1a
27,000 years ago
Around the Alps, and Germanic countries
J1b
23,000 years ago
Russia, South-East Europe, France, Italy and Iberia
J1b1
Britain, Ireland and Scandinavia
Germanic
J2
France, Italy and South-East Europe
J2a
19,000 years ago
Most of Europe
T*
10,000 years ago
Eastern Baltic, France, Italy
T1
Southern and Eastern Europe
T2
Northern, Central and Eastern Europe
Slavic-Germanic
T3
Iceland
Germanic
T4
Northern, Central and Eastern Europe
Germanic
T5
Central Europe and Britain
Celtic
Sources
The list below is non-exhaustive and include many of the numerous references linked on these websites. Some studies and databases not published on the Web were also used.
