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Haplogroup R and the Kura araxes culture

Two important subclades of Haplogroup R1a1a appear to broadly divide the European and Asian parts of this large clade: R-Z283 (R1a1a1b1) appears to encompass most of the R1a1a found in Europe (Pamjav 2012), while R-Z93 (R1a1a1b2) appears to encompass most of the R1a1a found in Asia.

These two subclades descends from Haplogroup R1a1a (M17)/(M198), which makes up the vast majority of all R1a over its entire geographic range. R1a1a descends it self from Haplogroup R1a1 (SRY1532.2), that originated as a single mutation of one male, the R1a1 originator considered to be the ancestor of all individuals carrying R1a1.

Haplogroup R1a1 descends from R1a (M420), which descends from Haplogroup R1 (M173), which it self is descends of Haplogroup R (M207). Haplogroup R1* and R2* might have originated in southern Central Asia (between the Caspian depression and the Hindu Kush).

Haplogroup R

Haplogroup R (M207) is believed to have arisen around 20,000-34,000 years ago (Karafet 2008), somewhere in Central Asia or South Asia, where its ancestor Haplogroup P-M45 is most often found at polymorphic frequencies (Wells 2001). It has been proven by ancient DNA to be at least 23,000 years.

Haplogroup R is found throughout all continents, but is fairly common throughout Europe, South Asia and Central Asia. Small frequencies are found in Malaysia, Indonesia, Philippines, and Indigenous Australians (Kayser 2003). It also occurs in Caucasus, Near East, West China, Siberia and some parts of Africa.

The two currently defined subclades are R1 (M173) and R2 (M479). Haplogroup R1 have a possible time of origin for 26,000 years ago while haplogroup R2 is estimated to be 12,000 years old.

The origins of R1 remain unclear. Haplogroup R is part of the family of haplogroup P (M45), and a sibling clade, therefore, of haplogroup Q (M242), which is common in the Americas and Eurasia. In Eurasia, Q’s geography includes eastern areas such as Siberia.

Based on these ancestral lineages, an inferred origin for R1 to the east of the West Asia. For example, Kivisild 2003 believes the evidence “suggests that southern and western Asia might be the source of this haplogroup” and “Given the geographic spread and STR diversities of sister clades R1 and R2, the latter of which is restricted to India, Pakistan, Iran, and southern central Asia, it is possible that southern and western Asia were the source for R1 and R1a differentiation.”

Soares 2010 felt in their review of the literature, that the case for South Asian origins is strongest, with the Central Asian origin argued by (Wells 2001) being also worthy of consideration.

Haplogroup R2 is defined by the presence of the marker M479. The paragroup for the R2 lineage is found in Pakistan, Portugal, Spain, Italy, and among Tatars in Bashkortostan in Russia and Ossetins in North Caucasus.

Haplogroup R1a (M420) probably branched off from R1* around the time of the Last Glacial Maximum 19,000 to 26,000 years ago. Little is know for certain about its place of origin. Some think it might have originated in the Balkans or around Pakistan and Northwest India, due to the greater genetic diversity found in these regions.

R1a is present at high frequency of 40 % plus from the Czech Republic across to the Altai Mountains in Siberia and south throughout Central Asia. Absolute dating methods suggest that this marker is 10–15,000 years old, and the microsatellite diversity is greatest in southern Asia.

The diversity can be explained by other factors though. The Balkans have been subject to 5000 years of migrations from the Eurasian Steppes, each bringing new varieties of R1a. South Asia has had a much bigger population than any other parts of the world (occasionally equalled by China) for at least 10,000 years, and larger population bring about more genetic diversity. The most likely place of origin of R1a is Central Asia or southern Russia/Siberia.

Haplogroup R1a1 (SRY1532.2) is defined by SRY1532.2, also referred to as SRY10831.2. SNP mutations understood to be always occurring with SRY1532.2 include SRY10831.2, M448, L122, M459, and M516.

This family of lineages is dominated by the R-M17 branch, which is positive for M17 and M198. The paragroup R-SRY1532.2* is positive for the SRY1532.2 marker but lacks either the M17 or M198 markers.

The R-SRY1532.2* paragroup is apparently less rare than R1* but still relatively unusual, though it has been tested in more than one survey.

Underhill 2009 for example report 1/51 in Norway, 3/305 in Sweden, 1/57 Greek Macedonians, 1/150 Iranians, 2/734 Ethnic Armenians, and 1/141 Kabardians.(Underhill 2009) While Sahoo 2006 reported R-SRY1532.2* for 1/15 Himachal Pradesh Rajput samples (Sahoo 2006).

Haplogroup R1a1a (M17)/(M198) makes up the vast majority of all R1a over its entire geographic range. It is defined by SNP mutations M17 or M198, which have always appeared together in the same men so far.

Two important subclades of haplogroup R1a1a appear to broadly divide the European and Asian parts of this large clade: R-Z283 (R1a1a1b1) appears to encompass most of the R1a1a found in Europe (Pamjav 2012), while R-Z93 (R1a1a1b2) appears to encompass most of the R1a1a found in Asia.

The modern distribution of R-M17 is distinctive. There are two widely separated areas of high frequency, one in South Asia, around India, and the other in Eastern Europe, around Poland and Ukraine. The demographic reasons for this are the subject of on-going discussion and attention among population geneticists and genetic genealogists, however, such patterns could be the combined result of (i) migrations and admixture, (ii) natural selection, and (iii) random genetic drift.

Despite deserved criticism by most archaeologists and anthropologists, even prominent historians and archaeologists have recently attempted to “marry” the evidence from the social sciences with that of genetic anthropology. Whilst the notion that genes, language and culture are co-eval is highly questionable, the link between R1a and “Indo-Europeans” remains a topic of considerable scholarly interest.

Until 2012, there was extensive scholarly debate as to the origins of haplgroup R-M17. This was a result of (i) a lack of further phylogenetic resolution of R-M17 into ‘daughter’ sub-clades and (ii) the evidently erroneous belief that measure of “STR diversity” can unambigiuosly qualify as to which population harbours the ‘oldest’ R-M17 haplogroups.

A large corpus of scholars had found that Indian, or more generally, South Asian populations, had the highest STR diversity. On the basis of these studies, and using the Evolutionary Effective Mutation Rate, several of the above authors concluded that R-M17 has been present in South Asian populations since the Neolithic, having originated there.

They further used this evidence to refute the hypothesis that R-M17 arrived with Indo-European invaders from the north. However, the use of this mutation rate has received criticism, as it should not be used with haplogroup populations which clearly show evidence of population expansion, such as R-M17.

Thus, using this mutation rate could artificially ‘blow out’ the actual age of R-M17 by as much as three-fold. Indeed, authors using the contrary, “germline mutation rate” (which is the rate empirically observed in father-son studies) arrive at more recent age estimates. In fact, Busby et al recently argued that the use of STR diversity in calculating ‘ages’ of haplogroups is highly problematical.

Other studies variously proposed Eastern European, Central Asian and even Western Asian origins for R – M17.

The decade-long debate as to which Eurasian region possessed the most diverse, hence oldest, STR values within R-M17 has been effectively put to an end with the discovery of R-M17 sub-clades. SNPs offer a clearer and more robust resolution than STRs. These findings have actually been known for a few years by genealogical companies and enthusiast genealogists, however, two academic, peer-reviewed papers were finally produced by Pamjav et al (2012, 2013). They discovered that all their tested Indian R-M17 samples belong to the Z-93 sub-clade, which is a derivative, “daughter” branch of R-M17.

In contrast, Eastern European populations belong to different daughter branches of R-M17, namely Z- 280 and M-458. The former is widely distributed over south-eastern, central-eastern and eastern Europe, and as far as Central Asia.(Pamjav 2012) Indeed, Central Asia “is an overlap zone for the R1a1-Z280 and R1a1-Z93”, being found in Mongol and Uzbek populations . On the other hand, M-458 is more geographically restricted to central-eastern Europe.

Furthermore, this study found that the undifferentiated, ‘parental’ M-198 existed in the European populations, but was not found in the Indian groups sampled (consisting of 256 Malaysian Indians, 301 Roma, 203 Dravidians from India).

Nevertheless, the authors concluded that “This pattern implies that an early differentiation zone of R1a1-M198 conceivably occurred somewhere within the Eurasian Steppes or the Middle East and Caucasus region as they lie between South Asia and Eastern Europe”, from where “South Asian’ Z-93 and “European” Z-283 sub-clades differentiated and spread in opposite directions.

Archaeologists recognize a complex of inter-related and relatively mobile cultures living on the Eurasian steppe, part of which protrudes into Europe as far west as Ukraine. These cultures from the late Neolithic and into the Iron Age, with specific traits such as Kurgan burials and horse domestication, have been associated with the dispersal of Indo-European languages across Eurasia.

Nearly all samples from Bronze and Iron Age graves in the Krasnoyarsk area in south Siberia belonged to R-M17 and appeared to represent an eastward migration from Europe.

In central Europe, Corded Ware period human remains at Eulau from which Y-DNA was extracted appear to be R-M17(xM458) (which they found most similar to the modern German R-M17* haplotype.

Origin

Haplogroup R1b (Y-DNA) is the dominant paternal lineage of Western Europe. In human genetics, Haplogroup R1b is the most frequently occurring Y-chromosome haplogroup in Western Europe and in parts of sub-Saharan Central Africa (for example around Chad and Cameroon). R1b is also present at lower frequencies throughout Eastern Europe, Western Asia, Central Asia, and parts of North Africa, South Asia, and Siberia.

Due to European emigration it also reaches high frequencies in the Americas and Australia. While Western Europe is dominated by the R1b1a2 (R-M269) branch of R1b, the Chadic-speaking area in Africa is dominated by the branch known as R1b1c (R-V88). These represent two very successful “twigs” on a much bigger “family tree.”

Early research focused upon Europe. In 2000 Ornella Semino and colleagues argued that R1b had been in Europe before the end of Ice Age, and had spread north from an Iberian refuge after the Last Glacial Maximum. Age estimates of R1b in Europe have steadily decreased in more recent studies, at least concerning the majority of R1b, with more recent studies suggesting a Neolithic age or younger.

Only Morelli et al. have recently attempted to defend a Palaeolithic origin for R1b1b2. Irrespective of STR coalescence calculations, Chikhi et al. pointed out that the timing of molecular divergences does not coincide with population splits; the TMRCA of haplogroup R1b (whether in the Palaeolithic or Neolithic) dates to its point of origin somewhere in Eurasia, and not its arrival in western Europe.

Barbara Arredi and colleagues were the first to point out that the distribution of R1b STR variance in Europe forms a cline from east to west, which is more consistent with an entry into Europe from Western Asia with the spread of farming. A 2009 paper by Chiaroni et al. added to this perspective by using R1b as an example of a wave haplogroup distribution, in this case from east to west.

The proposal of a southeastern origin of R1b were supported by three detailed studies based on large datasets published in 2010. These detected that the earliest subclades of R1b are found in western Asia and the most recent in western Europe.

While age estimates in these articles are all more recent than the Last Glacial Maximum, all mention the Neolithic, when farming was introduced to Europe from the Middle East as a possible candidate period. Myres et al. (August 2010), and Cruciani et al. (August 2010) both remained undecided on the exact dating of the migration or migrations responsible for this distribution, not ruling out migrations as early as the Mesolithic or as late as Hallstatt but more probably Late Neolithic.

They noted that direct evidence from ancient DNA may be needed to resolve these gene flows. Lee et al. (May 2012) analysed the ancient DNA of human remains from the Late Neolithic Bell Beaker site of Kromsdorf, Germany identifying two males as belonging to the Y haplogroup R1b. Analysis of ancient Y DNA from the remains of populations derived from early Neolithic settlements such as the Mediterranean Cardium and Central and North European LBK settlements have found an absence of males belonging to haplogroup R1b.

The Spread

The Dnieper River is one of the major rivers of Europe (fourth by length), rising near Smolensk and flowing through Russia, Belarus and Ukraine to the Black Sea. The total length is 2,145 kilometres (1,333 mi) with a drainage basin of 504,000 square kilometres (195,000 sq mi).

The river is noted for its dams and hydroelectric stations. The Dnieper is an important navigable waterway for the economy of Ukraine and is connected via the Dnieper-Bug Canal to other waterways in Europe.

In antiquity, the river was known to the Greeks as the Borysthenes and was part of the Amber Road. Arheimar, a capital of the Goths, was located on the Dnieper, according to the Hervarar saga.

Genetic studies based on modern male Scandinavian DNA suggest the advent of Ahrensburg “culture” may be associated with the bearers of the Haplogroup R1a1, that are postulated to have originally expanded to Europe and brought the reindeer hunters of the Ahrensburg “culture” from the Dniepr-Don Valley in Ukraine during the Late Glacial Maximum and reached Scandinavia between 13,000 and 7600 years ago.

Archaeological evidences have suggested that the core of the oldest populations colonising Scandinavia 11 00012 000 thousand years ago came from the present Germany. They probably went through the Jutland and the now submerged Doggerland, and then headed northward via the ice-free coastal area of Norway.

Their lithic tools suggest they belonged to the Ahrensburgian culture,that thrived in central and eastern Europe (artifacts have been found in Poland, Lithuania and Russia) in the so called ‘Late Glacial’, that is at the end of the LGM, when the increase of temperature and precipitation triggered the recession of the ice sheets.

The analysis of many Ahrensburg sites and the related lithic tools has suggested that this culture started from the Dniepr valley in Ukraine, one of the sites were humans found refuge during the LGM.

The analysis of Y chromosome polymorphisms in present European populations has indicated that Eu 19 (that is also characterised by other Y chromosome markers: 49a,fht11, SRY 1532G) expanded between 13 000 and 7600 years ago from the Dniepr-Don Valley area, probably when groups that initially sought refuge in that area during the LGM were allowed to migrate by the improved climatic conditions to those regions of Europe previously covered by ice.

In fact this Y chromosome lineage, is by far more frequent is eastern Europe with a decreasing westward gradient. In addition it is much more diversified in eastern European populations. The highest degree of diversification was observed in Ukranians.

It is then possible that Ahrensburgian men, as well as most of the men descending from the Ukranian LGM refuge bore Eu19 Y chromosomes. The microsatellite haplotypes linked to M17 in Norwegian individuals represents indeed a subset of the repertoire observed in eastern Europe. In particular it was observed the prevalence of the 15.3/1 (21/19 repeats) and of the 16.5/1 (23/19 repeats) haplotypes with their relative derivatives.

The Eu19 16.5/1 haplotype is also very common in eastern Europe, while Eu 19 15.3/1 haplotype is common in Norway, but very rare elsewhere. This peculiar pattern of microsatellites affiliated with EU19 may be explained by a founder effect, subsequent isolation in Norwegians (and possibly the Scandinavians) and eventual in loco expansion, as also observed elsewhere.

If it seems reasonable to assume that most of the Ahrensburgian men bore the Eu19 Y chromosomes, it cannot be excluded that they mixed with other groups before moving northward to Norwegian coasts. In particular, late glacial central Europe was characterised by the expansion of northern Balkan groups, where the frequency of M170 Y chromosomes (EU 7) was probably very high.

In addition, based on the differentiation of haplogroup V in Scandinavia, it also seems that groups coming from the northern Spain refuge entered Norway. Should this be true, it is likely that M173 Y chromosomes (EU18) also entered Norway during the late glacial.

R1a is also thought to have been the dominant haplogroup among the northern and eastern Proto-Indo-European language speakers, that evolved into the Indo-Iranian, Thracian, Baltic and Slavic branches. The Proto-Indo-Europeans originated in the Yamna culture (3300-2500 BCE).

With the exception of rather recent migrants from the Urals, it has been proposed that the appearance of domestic economies in Scandinavia arrived from central and northern Europe.

These populations would have likely been Indo-European speakers that possibly fostered the Proto Baltic-Slavic-Germanic linguistic unity to the Baltic area and to north-eastern Europe. Specifically the Corded Wares culture from Central Europe (present Germany) and the Battle-Axe culture from Jutland.

The spread of agriculture correlated with the Corded WaresBattle Axe cultures and possibly involved the displacement of some of the previous populations, but in other cases, such as Poland and northern Russia it was mainly a cultural phenomenon. In Norway, hunting and fishing became a secondary source of sustenance. However it is debated if this was a consequence of the displacement of the previous populations, or of a cultural switch driven by a few newcomers.

The present German gene pool shows a high frequency of Eu7 and Eu18 haplotypes. These haplotypes, which account for about 75% of the Norwegian Y chromosome pool, are then likely to have been brought to Norway by those groups who also brought the Indo-European languages and the agriculture.

However, at present, it is not possible to evaluate how much this migration impacted the Norwegian gene pool. First, because it is not possible to distinguish between lineages brought in the late glacial time and those brought 56000 years ago.

In addition, non-random mating phenomena may also play a role. It is possible that these cultures were composed of a subset of elite males, who reduced the reproductive success of other males and then the Y chromosome may emphasise the real genetic contributions of the central European migration to Norway.

It has to be noticed that also part of the Eu19 lineage could have entered Norway with the migrations that brought the transition to agriculture. In fact, although very low in Germany, this lineage is very high in Poland, Hungary and in the former Czeckslovakia. However the network of microsatellite haplotypes attached to this lineage seems to suggest a long time in loco differentiation.

Either because of late glacial or of more recent migrations the Norway Y chromosome gene pool appears to be very close to present day Germans. In fact the st and the Fst data indicate Germans and a few other Central European populations as being the closest to the Norwegians.

When we compare our results with those based on different polymorphic systems, we can infer that these conclusions are also valid for Swedish, while Finns and Saami had a quite different genetic history with a great impact of Uralic Finno-Ugric speaking population.

Ornella Semino et al. propose a postglacial spread of the R1a1 gene from the Ukrainian LGM refuge, subsequently magnified by the expansion of the Kurgan culture into Europe and eastward. The greatest variation in R1a1a is found in South Asia particularly North India. This conclusively proves that South Asia is the most probable source of R1a1a. Wells suggests the origin, distribution and age of R1a1 points to an ancient migration, possibly corresponding to the spread by the Kurgan people in their expansion across the Eurasian steppe around 3000 BC.

Their dramatic expansion was possible thanks to an early adoption of bronze weapons and the domestication of the horse in the Eurasian steppes (circa 4000-3500 BCE). The southern Steppe culture is believed to have carried predominantly R1b (M269 and M73) lineages, while the northern forest-steppe culture would have been essentially R1a-dominant.

The first expansion of the forest-steppe people occured with the Corded Ware Culture (see Germanic branch below). The migration of the R1b people to central and Western Europe left a vacuum for R1a people in the southern steppe around the time of the Catacomb culture (2800-2200 BCE).

The forest-steppe origin of this culture is obvious from the introduction of corded pottery and the abundant use of polished battle axes, the two most prominent features of the Corded Ware culture. This is also probably when the satemisation process of the Indo-European languages began since the Balto-Slavic and Indo-Iranian language groups belong to the same Satem isogloss and both appear to have evolved from the the Catacomb culture.

Ancient DNA testing has confirmed the presence of haplogroup R1a1a in samples from the Corded Ware culture in Germany (2600 BCE), from Tocharian mummies (2000 BCE) in Northwest China, from Kurgan burials (circa 1600 BCE) from the Andronovo culture in southern Russia and southern Siberia, as well as from a variety of Iron-age sites from Russia, Siberia, Mongolia and Central Asia.

Across Caucasus

The Tell Halaf/Ubaid culture are tied both to the Southern Levant and then to the Trancausia area and possibly inspiring both Kura-Axes and Maykop development. There are some very early branches of Haplogroup R1b1a2 (M269) in the Near East, as well as the non M269 R1b1c (V88) that may have expanded into Africa from there. The Kura-Axes and Maykop area becomes core to the Circumpontic Metallugy Province. We have M269 L23xL51 in the Caucasus and Anatolia and L51 types of R1b showing up with Bell Beakers, metallurgists, in Western Euroope. 

The Paleolithic origins of R1b are not entirely clear to this day. Some of the oldest forms of R1b are found around the Caucasus, in Iran and in southern Central Asia, a vast region where could have roamed the nomadic R1b hunter-gatherers during the Ice Age. A branch of R1 would have developed into R1b then R1b1 and R1b1a in the northern part of the Middle East around the time of the Last Glacial Maximum (circa 20,000 years ago), while R1a migrated north to Siberia.

R1b1a presumptively moved to northern Anatolia and across the Caucasus during the Neolithic, where it split into R1b1a1 (M73) and R1b1a2 (M269). The Near Eastern leftovers evolved into R1b1c (V88), now found at low frequencies among the Lebanese, the Druze, and the Jews. The Phoenicians (who came from modern day Lebanon) spread this R1b1c to their colonies, notably Sardinia and the Maghreb.

R1b1a2 (the most common form in Europe) and R1b1a1 is closely associated with the diffusion of Indo-European languages, as attested by its presence in all regions of the world where Indo-European languages were spoken in ancient times, from the Atlantic coast of Europe to the Indian subcontinent, including almost all Europe (except Finland and Bosnia-Herzegovina), Anatolia, Armenia, European Russia, southern Siberia, many pockets around Central Asia (notably Xinjiang, Turkmenistan, Tajikistan and Afghanistan), without forgetting Iran, Pakistan, India and Nepal. The history of R1b and R1a are intricately connected to each others.

Modern linguists have placed the Proto-Indo-European homeland in the Pontic-Caspian Steppe, a distinct geographic and archeological region extending from the Danube estuary to the Ural mountains to the east and North Caucasus to the south. The Neolithic, Eneolithic and early Bronze Age cultures in Pontic-Caspian steppe has been called the Kurgan culture (7000-2200 BCE) by Marija Gimbutas, due to the lasting practice of burying the deads under mounds (“kurgan”) among the succession of cultures in that region. It is now known that kurgan-type burials only date from the 4th millenium BCE and almost certainly originated south of the Caucasus.

Horses were first domesticated around 4600 BCE in the Caspian Steppe, perhaps somewhere around the Don or the lower Volga, and soon became a defining element of steppe culture. Nevertheless it is unlikely that R1b was already present in the eastern steppes at the time, so the domestication of the horse should be attributed to the indigenous R1a people.

It is not yet entirely clear when R1b crossed over from eastern Anatolia to the Pontic-Caspian steppe. This could have happened during or just after the Neolithic, or both. The genetic diversity of R1b being greater around the Caucasus it is hard to deny that R1b evolved there before entering the steppe world.

It is possible that a first R1b migration from Anatolia in the 5th or even 6th millennium BCE introduced sheep into the steppe, an animal whose wool would play an important role in Celtic and Germanic (R1b branches of the Indo-Europeans) clothing traditions up to this day. Another migration across the Caucasus happened shortly before 3700 BCE, when the Maykop culture, the world’s first Bronze Age society, appeared apparently out of nowhere in the north-west Caucasus.

The origins of Maykop are still uncertain, but archeologists have linked it to contemporary Chalcolithic cultures in Assyria and western Iran. Archeology also shows a clear diffusion of bronze working and kurgan-type burials from the Maykop culture to the Pontic Steppe, where the Yamna culture (3500-2500 BCE) developed soon afterwards (from 3500 BCE).

Middle Eastern R1b people had been living and blending to some extent with the local R1a foragers and herders for over a millennium, perhaps even two or three. The close cultural contact and interactions between R1a and R1b people all over the Pontic-Caspian Steppe resulted in the creation of a common vernacular, a new lingua franca, which linguists have called Proto-Indo-European (PIE).

Linguistic similarities exist between PIE and Caucasian and Hurrian languages in the Middle East on the one hand, and Uralic languages in the Volga-Ural region on the other hand, which makes the Pontic Steppe the perfect intermediary region. Kurgan (a.k.a. tumulus) burials would become a dominant feature of ancient Indo-European societies and were widely used by the Celts, Romans, Germanic tribes, and Scythians, among others.

During the Yamna period cattle and sheep herders adopted wagons to transport their food and tents, which allowed them to move deeper into the steppe, giving rise to a new mobile lifestyle that would eventually lead to the great Indo-European migrations. This type of mass migration in which whole tribes moved with the help of wagons was still common in Gaul at the time of Julius Caesar, and among Germanic peoples in the late Antiquity.

The Yamna horizon was not a single, unified culture. In the south, along the northern shores of the Black Sea coast until the the north-west Caucasus, was a region of open steppe, expanding eastward until the Caspian Sea, Siberia and Mongolia (the Eurasian Steppe).

The western section, between the Don and Dniester Rivers (and later the Danube), was the one most densely settled by R1b people, with only a minority of R1a people (5-10%). The eastern section, in the Volga basin until the Ural mountains, was inhabited by R1a people with a substantial minority of R1b people (whose descendants can be found among the Bashkirs, Turkmans, Uyghurs and Hazaras, among others). The northern part of the Yamna horizon was forest-steppe occupied by R1a people, also joined by a small minority of R1b (judging from modern Russians and Belarussians, the frequency of R1b was from seven to nine times less lower than R1a).

The western branch would migrate to the Balkans and Greece, then to central and Western Europe, and back to their ancestral Anatolia in successive waves (Hittites, Phrygians, Armenians, etc.). The eastern branch would migrate to Central Asia, Xinjiang, Siberia, and South Asia (Iran, Pakistan, India). The northern branch would evolve into the Corded Ware culture and disperse around the Baltic, Poland, Germany and Scandinavia.

Maykop was an advanced Bronze Age culture, actually one of the very first to develop metalworking, and therefore metal weapons. The world’s oldest sword was found at a late Maykop grave in Klady kurgan 31. Its style is reminiscent of the long Celtic swords, though less elaborated.

Horse bones and depictions of horses already appear in early Maykop graves, suggesting that the Maykop culture might have been founded by steppe people or by people who had close link with them. However, the presence of cultural elements radically different from the steppe culture in some sites could mean that Maykop had a hybrid population.

Without DNA testing it is impossible to say if these two populations were an Anatolian R1b group and a G2a Caucasian group, or whether R1a people had settled there too. The two or three ethnicities might even have cohabited side by side in different settlements.

The one typical Caucasian Y-DNA lineage that does follow the pattern of Indo-European migrations is G2a3b1, which is found throughout Europe, Central Asia and South Asia. In the Balkans, the Danube basin and Central Europe its frequency is somewhat proportional to the percentage of R1b.

Maykop people are the ones credited for the introduction of primitive wheeled vehicles (wagons) from Mesopotamia to the steppes. This would revolutionise the way of life in the steppe, and would later lead to the development of (horse-drawn) war chariots 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, the western branch (R1b) of the Indo-Europeans from the Black Sea shores are excellent candidates for being the mysterious Sea Peoples, who raided the eastern shores of the Mediterranean during the second millennium BCE.

The rise of the IE-speaking Hittites in Central Anatolia happened a few centuries after the disappearance of the Maykop and Yamna cultures. Considering that most Indo-European forms of R1b found in Anatolia today belong to the R1b-Z2103 subclade, it makes little doubt that the Hittites came to Anatolia via the Balkans, after Yamna/Maykop people invaded Southeast Europe.

The Maykop and Yamna cultures were succeeded by the Srubna culture (1600-1200 BCE), possibly representing an advance of R1a1a people from the northern steppes towards the Black Sea shores, filling the vacuum left by the R1b tribes who migrated to Southeast Europe and Anatolia.

There is substantial archaeological and linguistic evidence that Troy was an Indo-European city associated with the steppe culture and haplogroup R1b. The Trojans were Luwian speakers related to the Hittites (hence Indo-European), with attested cultural ties to the culture of the Pontic-Caspian steppe.

The Hittites (c. 2000-1178 BCE) were the first Indo-Europeans to defy (and defeat) the mighty Mesopotamian and Egyptian empires. There are two hypotheses regarding the origins of the Hittites. The first is that they came from the eastern Balkans and invaded Anatolia by crossing the Bosphorus. That would mean that they belonged either to the L23 or the Z2103 subclade.

The other plausible scenario is that they were an offshoot of the late Maykop culture, and that they crossed the Caucasus to conquer the Hattian kingdom (perhaps after being displaced from the North Caucasus by the R1a people of the Catacomb culture). In that case the Hittites might have belonged to the R1b-M269 subclade.

The first hypothesis has the advantage of having a single nucleus, the Balkans, as the post-Yamna expansion of all Indo-European R1b. The Maykop hypothesis, on the other hand, would explain why the Anatolian branch of IE languages (Hittite, Luwian, Lydian, Palaic) is so archaic compared to other Indo-European languages, which would have originated in Yamna rather than Maykop.

The first city of Troy dates back to 3000 BCE, right in the middle of the Maykop period. Troy might have been founded by Maykop people as a colony securing the trade routes between the Black Sea and the Aegean. The founding of Troy happens to coincide exactly with the time the first galleys were made. Considering the early foundation of Troy, the most likely of the two Indo-European paternal haplogroups would be R1b-M269 or L23.

R1b1a2

R1b1a2 (2011 name) is defined by the presence of SNP marker M269. R1b1a2* or M269(xL23) is found at highest frequency in the central Balkans notably Kosovo with 7.9%, Macedonia 5.1% and Serbia 4.4%. Kosovo is notable in also having a high percentage of descendant L23* or L23(xM412) at 11.4% unlike most other areas with significant percentages of M269* and L23* except for Poland with 2.4% and 9.5% and the Bashkirs of southeast Bashkortostan with 2.4% and 32.2% respectively. Notably this Bashkir population also has a high percentage of M269 sister branch M73 at 23.4%. Five individuals out of 110 tested in the Ararat Valley, Armenia belonged to R1b1a2* and 36 to L23*, with none belonging to subclades of L23.

European R1b is dominated by R-M269. It has been found at generally low frequencies throughout central Eurasia, but with relatively high frequency among Bashkirs of the Perm Region (84.0%). This marker is also present in China and India at frequencies of less than one percent. The table below lists in more detail the frequencies of M269 in various regions in Asia, Europe, and Africa.

The frequency is about 71% in Scotland, 70% in Spain and 60% in France. In south-eastern England the frequency of this clade is about 70%; in parts of the rest of north and western England, Spain, Portugal, Wales and Ireland, it is as high as 90%; and in parts of north-western Ireland it reaches 98%. It is also found in North Africa, where its frequency surpasses 10% in some parts of Algeria.

As discussed above, in articles published around 2000 it was proposed that this clade been in Europe before the last Ice Age, but by 2010 more recent periods such as the European Neolithic have become the focus of proposals.

A range of newer estimates for R1b1b2, or at least its dominant parts in Europe, are from 4,000 to a maximum of about 10,000 years ago, and looking in more detail is seen as suggesting a migration from Western Asia via southeastern Europe. Western European R1b is dominated by R-P310.

It was also in this period between 2000 and 2010 that it became clear that especially Western European R1b is dominated by specific sub-clades of R-M269 (with some small amounts of other types found in areas such as Sardinia).

Within Europe, R-M269 is dominated by R-M412, also known as R-L51, which according to Myres et al. (2010) is “virtually absent in the Near East, the Caucasus and West Asia.” This Western European population is further divided between R-P312/S116 and R-U106/S21, which appear to spread from the western and eastern Rhine river basin respectively.

Myres et al. note further that concerning its closest relatives, in R-L23*, that it is “instructive” that these are often more than 10% of the population in the Caucasus, Turkey, and some southeast European and circum-Uralic populations. In Western Europe it is also present but in generally much lower levels apart from “an instance of 27% in Switzerland’s Upper Rhone Valley.”

In addition, the sub-clade distribution map, Figure 1h titled “L11(xU106,S116)”, in Myres et al. shows that R-P310/L11* (or as yet undefined subclades of R-P310/L11) occurs only in frequencies greater than 10% in Central England with surrounding areas of England and Wales having lower frequencies.

This R-P310/L11* is almost non-existent in the rest of Eurasia and North Africa with the exception of coastal lands fringing the western and southern Baltic (reaching 10% in Eastern Denmark and 6% in northern Poland) and in Eastern Switzerland and surrounds.

In 2009, DNA extracted from the femur bones of 6 skeletons in an early-medieval burial place in Ergolding (Bavaria, Germany) dated to around 670 AD yielded the following results: 4 were found to be haplogroup R1b with the closest matches in modern populations of Germany, Ireland and the USA while 2 were in Haplogroup G2a.

Population studies which test for M269 have become more common in recent years, while in earlier studies men in this haplogroup are only visible in the data by extrapolation of what is likely. The following gives a summary of most of the studies which specifically tested for M269, showing its distribution in Europe, North Africa, the Middle East and Central Asia as far as China and Nepal.

Armenia

The geographical distribution of this haplotype is such that it is shared by Armenians and two other populations from the Caucasus. Moreover, it is lacking in most other populations from the Caucasus, as well as in the other populations from further east. On the other hand, it is more frequently found in Europe, where as we know, haplogroup R1b tends to have higher frequencies as well.

The Armenian modal haplotype is also the modal R1b3 haplotype observed by Cinnioglu in Anatolia. According to him, apparently it entered Anatolia from Europe in Paleolithic times, and diffused again from Anatolia in the Late Upper Paleolithic.

An alternative explanation may be that the particular haplotype may have been associated with the movement of the Phrygians into Asia Minor. The Phrygians were an Indo-European people of the Balkans who settled in Asia Minor, and the Armenians were reputed to be descended from them.

It would be interesting to thoroughly study the populations of modern Thrace, Anatolia, and Armenia, and to investigate whether a subgroup of R1b3 chromosomes linked by the Armenian modal haplotype may represent the signature of a back-migration into Asia of Balkan Indo-European peoples.

Combined with advanced bronze weapons and their sea-based culture, the western branch (R1b) of the Indo-Europeans from the Black Sea shores are excellent candidates for being the mysterious Sea Peoples, who raided the eastern shores of the Mediterranean during the second millennium BCE.

The Phrygians and the Proto-Armenians are two other Indo-European tribes stemming from the Balkans. Both appear to have migrated to Anatolia around 1200 BCE, during the ‘great upheavals’ of the Eastern Mediterranean.

The Phrygians (or Bryges) founded a kingdom (1200-700 BCE) in west central Anatolia, taking over most of the crumbling Hittite Empire. The Armenians crossed all Anatolia until Lake Van and settled in the Armenian Highlands. Nowadays 30% of Armenian belong to haplogroup R1b, the vast majority to the L23 subclade.

Armenian DNA Project – News

Kura-Araxes culture

The period between the 4th and 3rd millennia B.C. was the time of great cataclysmic events in the Caucasus; its cultural advances were influenced by changes within its boundaries as well as interactions with the outside world. The most significant occurrence of this epoch was the appearance of a large number of peoples of Mesopotamian cultural identity who contributed to speeding up the rhythm of its cultural development, adding “explosive” character to its progress.

During this period the South Caucasus experienced two powerful waves of Middle Eastern expansion: the first at the time of Late Neolithic culture of Sioni in the 4th-5th millennia B.C., and the second at the period of Tsopi culture in the Late Neolithic Age, at the end of the 5th and the first half of the 4th millennium B.C., which is known as the Uruk expansion era.

Later, in the second half of the 4th and throughout the 3 rd millennium B.C., during the Early Bronze Age the Kura-Araxes culture of the Caucasus spread throughout the greater part of the Caucasus, Eastern Anatolia, northern parts of Iran, Middle East and even Europe.

In this context, recent archaeological finds in the Southern and Northeastern Caucasus gave yet another, entirely new nuance to scientific researches into the ancient past of the Caucasus. They made it clear that incursion of these peoples into the Caucasus was not a onetime event, but continued for a significantly long period.

Reasoning by the topography of the archaeological finds in Mesopotamia, it becomes clear that large masses of migrant settlers from that area did not move straight along the route to Transcaucasia in order to reach the destination faster. Actually, they settled down in every region of the Caucasus, in the mountains and flatlands, in areas where they could maintain a lifestyle familiar to them.

It seems obvious that from that period on, two cultures of the Caucasus that had been at different stages of development could coexist peacefully on the basis of their mutual participation in metallurgical manufacturing; it was this type of communal economy that gave impetus to a speedy development of the local culture. This is well illustrated by the metallurgical items of the Kura-Araxes culture, which is significantly more advanced in comparison with the preAeneolithic culture.

At present the situation has changed drastically. On the basis of a whole series of radiocarbon analyses, it has been proved that burial mounds of the ancient pit-grave culture are of a significantly later period in comparison with Maikop archaeological sites.

This allows scholars to assume that the tradition of building this type of burial mounds emerged precisely in the Maikop culture. Its ties with Levant and Mesopotamian antiquities point to its earlier origin. At the same time, a whole range of chronological data obtained with radiocarbon analysis has established that the settlements and burial mounds of the South Caucasus containing Uruk artefact are coexistent with the Maikop culture and, accordingly, the ancient pit-grave culture and its burial mounds belong to a later period.

Therefore, today we cannot possibly ascribe the emergence of this kind of burial mounds in the Maikop culture as well as similar contemporaneous sites in the South Caucasus to the influence of the steppe zone cultures. Moreover, there were no adverse conditions that would have prevented emergence of this type of burial mounds in the Caucasus itself

Akhundov (2007) recently uncovered pre-Kura-Araxes/Late Chalcolithic materials from the settlement of Boyuk Kesik and the kurgan necropolis of Soyuq Bulaq in northwestern Azerbaijan, and Makharadze (2007) has also excavated a pre-Kura-Araxes kurgan, Kavtiskhevi, in central Georgia.

Materials recovered from both these recent excavations can be related to remains from the metal-working Late Chalcolithic site of Leilatepe on the Karabakh steppe near Agdam (Narimanov et al. 2007) and from the earliest level at the multi-period site of Berikldeebi in Kvemo Kartli (Glonti and Dzavakhishvili 1987). They reveal the presence of early 4th millennium raised burial mounds or kurgans in the southern Caucasus.

Similarly, on the basis of her survey work in eastern Anatolia north of the Oriental Taurus mountains, C. Marro (2007)likens chafffaced wares collected at Hanago in the Sürmeli Plain and Astepe and Colpan in the eastern Lake Van district in northeastern Turkey with those found at the sites mentioned above and relates these to similar wares (Amuq E/F) found south of the Taurus Mountains in northern Mesopotamia.

The Kura–Araxes culture or the early trans-Caucasian culture was a civilization that existed from 3400 BC until about 2000 BC, which has traditionally been regarded as the date of its end, but it may have disappeared as early as 2600 or 2700 BC.

The name of the culture is derived from the Kura and Araxes river valleys. Its territory corresponds to parts of modern Armenia, Azerbaijan, Chechnya, Dagestan, Georgia, Ingushetia and North Ossetia.

The ‘homeland’ (itself a very problematic concept) of the Kura-Araxes culture-historical community is difficult to pinpoint precisely, a fact that may suggest that there is no single well-demarcated area of origin, but multiple interacting areas including northeastern Anatolia as far as the Erzurum area, the catchment area drained by the Upper Middle Kura and Araxes Rivers in Transcaucasia and the Caspian corridor and adjacent mountainous regions of northeastern Azerbaijan and southeastern Daghestan.

While broadly (and somewhat imprecisely) defined, these regions constitute on present evidence the original core area out of which the Kura-Araxes ‘culture-historical community’ emerged.

In other words, sometime around the middle of the 4th millennium BCE or slightly subsequent to the initial appearance of the Maikop culture of the NW Caucasus, settlements containing proto-Kura-Araxes or early Kura-Araxes materials first appear across a broad area that stretches from the Caspian littoral of the northeastern Caucasus in the north to the Erzurum region of the Anatolian Plateau in the west.

For simplicity’s sake these roughly simultaneous developments across this broad area will be considered as representing the beginnings of the Early Bronze Age or the initial stages of development of the KuraAraxes/Early Transca