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Bell Beaker expansion

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Steppe ancestry has been recently found widespread in central European Bell Beaker individuals, while in Iberian samples this ancestry was not found, therefore excluding a substantial contribution to central Europe from Iberian Beaker Complex-associated individuals[Olalde et al. 2017]. This further supports Heyd’s archaeological model[Harrison and Heyd 2007][Heyd 2012][Heyd 2014], and contradicts the previous model of population expansion out of Iberia.

genetics.jpg

Modified file from recent papers on ancient samples from Eastern European, Southeastern European, Western European, and Bell Beaker cultures: Left: ADMIXTURE clustering analysis with k=8 showing ancient individuals. E/M/MLN, Early/Middle/Middle Late Neolithic; W/E/S/CHG, Western/Eastern/Scandinavian/Caucasus hunter-gatherers[Olalde et al. 2017]. Center: Supervised ADMIXTURE plot, modeling each ancient individual (one per row), as a mixture of populations represented by clusters containing Anatolian Neolithic (grey), Yamnaya from Samara (orange), EHG (red) and WHG (blue). Dates indicate approximate range of individuals in each population[Mathieson et al. 2017]. Right: Ancestral components in ancient individuals estimated by ADMIXTURE (k=11)[Mittnik et al. 2017].

In the lower Danube clear patterns of patrilocality and female exogamy have been found, apart from a continuing kinship relation in the transition of the Chalcolithic to the Bronze Age. There is evidence of continuing traditions from the Bell Beaker cultures to Early Bronze Age cultures in the region, with female mobility as a force for regional and supraregional communication and exchange[Knipper et al. 2017].

olalde-steppe.png

Image from Olalde et al. (2017). «Proportion of Steppe-related ancestry (shown in black) in Beaker Complex groups, computed with qpAdm under the model Yamnaya_Samara + Anatolia_Neolithic + WHG. The area of the pie is proportional to the number of individuals (shown inside the pie if more than one)»”. Original images under a CC-BY-NC 4.0 International license.

Bell Beaker samples from Hungary at overlapping dates show very different proportions (from 0% to 74%) of steppe ancestry, a heterogeneity consistent with early stages of admixture between European farmers and Yamna migrants. Also, in southern France and Britain there is a greater affinity to the Iberian Early Neolithic farmers than to central European farmers, which suggests a previous spread of migrants along the Atlantic coast, consistent with the interaction of Megalithic-building cultures involving movements of people[Olalde et al. 2017].

Bell Beaker samples from Hungary at overlapping dates show very different proportions (from 0% to 74%) of steppe ancestry, a heterogeneity consistent with early stages of admixture between European farmers and Yamna migrants. Also, in southern France and Britain there is a greater affinity to the Iberian Early Neolithic farmers than to central European farmers, which suggests a previous spread of migrants along the Atlantic coast, consistent with the interaction of Megalithic-building cultures involving movements of people (Olalde et al. 2017).

olalde-pca.png

Image from Olalde et al. (2017). «Principal component analysis of 990 present-day West Eurasian individuals (grey dots), with previously published (pale yellow) and new ancient samples projected onto the first two principal components»”. Original images under a CC-BY-NC 4.0 International license.

The oldest samples from the Bell Beaker culture are two individuals from Kromsdorf dated ca. 2550 BC, one of R1b1a1a2-M269 (xR1b1a1a2a1a1-U106), and the other of R1b-M343 (M269 unclear) lineages[Lee et al. 2012]. The oldest samples of R1b1a1a2a1a2-P312 lineages are found in Osterhofen ca. 2540 BC, and in Sierentz – Les villas d’Aurèle ca. 2430 BC. The oldest sample of haplogroup R1b1a1a2a1a2b-U152 is found in Budapest ca. 2335 BC, of haplogroup R1b1a1a2a1a2a-DF27[1] in Quedlinburg ca. 2290 BC[Lazaridis et al. 2016], and of haplogroup R1b1a1a2a1a2c-L21 in Amesbury ca. 2290 BC[Olalde et al. 2017].

chalcolithic_late_Europe.jpg Diachronic map of migrations in Europe ca. 2600-2250 BC[Heyd 2014][Anthony 2007][Sjogren, Price, and Kristiansen 2016][Harrison and Heyd 2007][Prieto Martínez and Salanova 2015][Fokkens and Nicolis 2012].

Regarding the arrival of Bell Beakers into Britain – marked by the appearance of steppe ancestry in aDNA samples –, it has been found that they are more closely related to lower Rhine individuals, and not to the Atlantic façade of western Europe[Olalde et al. 2017]. Demic diffusion of R1b1a1a2a1a2c1-L21 lineages accompanying Bell Beaker expansion in the British Isles was already supported by ancient DNA analysis[Cassidy et al. 2016], and all samples of the British Isles have been reported as of R1b1a1a2a1a2c-L21 (or older haplogroups). These data and the modern distribution of L21 subclades associated with the British Isles point more likely to a single successful migration of clans of R1b1a1a2a1a2c-L21 lineages into Britain. The contended Pre-Celtic Irish and Pictish substrates might have been therefore of Late Indo-European nature, imported by peoples of R1b1a1a2a1a2c1-L21 lineages.

Recent research already supported a considerable degree of mobility with little difference between male and female migration in Britain[Parker Pearson et al. 2016], and not an exchange of female marriage partners[Brodie 2001] or inter-cultural contact consolidation[Vander Linden 2007], as previously proposed.

r1b-L21.jpg Modern distribution of haplogroup R1b1a1a2a1a2c1-L21. Adapted from Myres et al.[Myres et al. 2011]

Studies of ancient Indo-European hydronymy[Krahe 1964][Krahe 1949][Nicolaisen 1957] have revealed a quasi-uniform name-giving system for water courses that shows Indo-European water-words and suffixes following rules of Late Proto-Indo-European word formation[Adrados 1998]. This points to an ancient wave of Late Indo-European speakers spread over Western and Central Europe before the Celtic and Germanic expansions, including the British Isles, the Italian and Iberian peninsulas.

The four certain samples of R1b1a1a2a1a2b-U152 lineage have been found in Bell Beaker territories from east to west Europe. The presence of a potential R1b1a1a2a1a2a-DF27 lineage in the central Bell Beaker group, and its subclade distribution in the modern west European population might be linked to this lineage’s expansion to the west and south during the Bell Beaker phenomenon.

The expansion of R1b1a1a2a1a-L151 lineages could then be linked to the first introduction of Indo-European languages in Western Europe[Cassidy et al. 2016], which could have left some traces of their presence in historic times. However, the later expansion of Celtic languages, and an apparent resurgence of the probably indigenous Proto-Iberian and Proto-Basque languages – possibly the descendant of the languages of early farmers, similar to Paleo-Sardinian[Terradas et al. 2014] – over an expanding Iberian subclade (R1b1a1a2a1a2a1b1a1-M167/SRY2627) of the R1b1a1a2a1a2a-DF27 lineage[Gunther et al. 2015] have left scarce data on the older situation.

Samples from northern and south-western Iberia in the transition from the Chalcolithic to the Bronze Age have been found to have steppe ancestry, although in lesser proportion than in central and western Europe[Olalde et al. 2017][Martiniano et al. 2017]. Samples of north-east Iberia, dated ca. 2385 BC, show haplogroups R1b1a-L754 (xR1b1a1a2a-L23), G2-P287, and I2a2-M436, which point to at least a partial persistence of old European hunter-gatherer lineages in the region.

However, there is a clear replacement of autochthonous I2-M438 and G2a-P15 subclades by R1b1a1a2a1a2-P312. The expansion of haplogroup R1b1a1a2a1a2a-DF27 from northern Iberia, based on studies of the modern population, also points to ca. 2200 BC[Solé-Morata et al. 2017]. The greater admixture with the autochthonous population probably suggests a different type of expansion than that of Bell Beakers in central Europe and the British Isles, probably a later renewed male-driven invasion from northern Iberia, once admixed with the local population.

The only certain Indo-European language of Iberia that can be considered of a non-Celtic nature is Lusitanian (which has been linked to a potential Galaico-Lusitanian group of the north-western Iberian Peninsula), and there has been some discussion on the pre-Celtic nature of the languages of Cantabri, Astures, Pellendones, Carpetani, and Vettones. Also, while the position of Tartessian as Indo-European[Koch 2009] is highly doubted[2] – , there is some support for a borrowing of names from a “lost Indo-European language” over the course of long-term contacts[Mikhailova 2015].

The emergence of El Argar groups was preceded by a break in Chalcolithic cultural traditions in south-east Iberia, which points to an upheaval of existing social structures or an influx of groups that cannot be distinguished from the local population at the present of genetic resolution, e.g. from south-eastern Europe[Szecsenyi-Nagy et al. 2017]. This could point to the time of resurge of groups associated with previous Neolithic cultures that might have conserved Pre-Iberian and Pre-Basque languages until historic times.

The other region where modern R1b1a1a2a1a2a-DF27 lineages peak in the modern population corresponds to the old Nordwestblock cultural region, where a non-Celtic, non-Germanic Indo-European language might have been spoken[Kuhn, Hachmann, and Kossack 1986].

r1b-DF27.jpg Modern distribution of haplogroup R1b1a1a2a1a2a-DF27. Adapted from Myres et al.[Myres et al. 2011]

To the east, in the Vistula group of Lesser Poland, Bell Beaker samples of R1b1a1a2-M269 are found from 2400 BC to 2300 BC. Later an eastern sample is found near the Oder ca. 2170 BC, which – together with long-term and long-distance economic exchange (especially regarding amber imports) during the Bronze Age[Makarowicz 2009] – support the presence of Old European river names in east Europe dating to this period.

References

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Notes

  1. ZZ11+ equivalent (ancestral to DF27 and U152); DF27+? In 390k BAM file, but short, may actually belong to chromosomes 2 or 5. Additional information from Alex Williamson.
  2. It was criticized extensively in a special section of Vol. 42 of The Journal of Indo-European Studies (No. 3 & 4, Fall/Winter 2014)