Corded Ware migration
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The Corded Ware culture has been argued to practice exogamy – most adult women being of non-local origin – based on a recent work on diet and mobility[Sjogren, Price, and Kristiansen 2016], and mtDNA has been documented to be more varied among Corded Ware females than men[Lazaridis et al. 2014]. Exogamy is described as a well-established practice over a long period of time, and the origin of females has been linked to Neolithic cultures still residing on the higher elevations in the region shared with Corded Ware cultures that colonized rivers and valleys[Kristiansen et al. 2017].
A violent picture has been proposed for the Corded Ware culture society[Haak et al. 2008], with warrior youth bands with seasonal activities, in a way similar to that documented in the Russian steppe from the Bronze Age onwards[Kristiansen et al. 2017].
In a first stage of the Corded Ware migrations, their nomadic way of life is represented by site-camps close to tumuli, located on routes of their long-lasting travels. In a second phase, semi-nomadic economic allowed for settlement micro-regions to appear, and cemeteries were a kind of landmark during the population’s seasonal migrations[Machnik 2004]. Necropolises were significant areas to which they returned periodically, establishing eventually site-camps, beginning their transformation into Early Bronze Age groups with a more stable economy[Witkowska 2006].
Analysis of aDNA has revealed that the plague was a prehistoric disease, associated with the Eurasian steppes, and linked to the Corded Ware culture expansion[Rasmussen et al. 2015][Andrades Valtueña et al. 2017], which connected vast areas in central-east Europe in a relatively short period. Also, the population to the north of the loess belt of the northern European plain is known to have been much lesser than in south-east Europe[Müller 2013].
This demographic disadvantage of central-east European lands might have provided part of the ‘push’ for the migration and expansion of the Corded Ware population[Anthony and Brown 2017], which show a clear genetic homogeneity – close to the Yamna component – from western to eastern Europe, in spite of the proposed practice of exogamy. This explosive migration over a sparsely populated area accounts for the clear founder effect of R1a1a1b-Z645 lineages, possibly originally from the Dniester-Dnieper or Middle Dnieper region.
The most recent sample from Zvejnieki, of the Late Neolithic – Corded Ware (ca. 2885 BC), clusters quite closely with Yamna samples, revealing a break of the regional population with the samples from a thousand years earlier, which were closer to Eastern European hunger-gatherer ancestry[Mathieson et al. 2017]. This close cluster of the Baltic with east Yamna, that does not show more EHG admixture – as expected in a sample potentially generations away from southern migrations (and thus admixed with the local population) – could point to the previous presence of a component similar to the Yamna ancestry in the neighbouring region of east-central Europe.
R1a1a1b-Z645, split from R1a1a1-M417 ca. 3500 BC, shows a TMRCA of ca. 3000 BC, coinciding with the formation date for mainly-European subclades R1a1a1b1-Z283 and R1a1a1b1a-Z282, and mainly-Asian subclade R1a1a1b2-Z93. The common TMRCA for R1a1a1b1-Z283 and R1a1a1b1a-Z282 suggests an expansion at nearly the same time as peoples of Corded Ware cultures are supposed to have migrated east- and westward, reaching the Middle Elbe-Saale region, where most aDNA samples analysed come from, about 2750 BC. The common TMRCA of 2700 BC for modern Asian lineages gives support to a later successful expansion into Asia centred on the eastern part of the Pontic-Caspian steppes (see Indo-Iranian).
Y-DNA samples of haplogroup R1a-M420 (probably R1a1a1-M417) are found in central Corded Ware culture groups[Allentoft et al. 2015][Mathieson et al. 2015][Haak et al. 2008][Mittnik et al. 2017][Saag et al. 2017], with one sample in Bergrheinfeld (ca. 2647), two samples in Eulau (ca. 2600 BC), one sample from Kyndelöse (ca. 2670 BC), seven samples from Esperstedt (one dated ca. 2430 BC and other six ca. 2275 BC), and two from the Battle Axe culture, one from Viby (ca. 2500 BC), and one from Ölsund (ca. 2350 BC). Five samples from Erperstedt have been inferred to be relatives via paternal line, which confirms their common sharing of bottom subclade R1a1a1-M417[Monroy Kuhn, Jakobsson, and Günther 2017].
Two samples from Tiefbrunn of haplogroups R-M207 (ca. 2755 BC) and R1b1-L278 (ca. 2725 BC) are of dubious subclade – the first could be R1a1a1-M417 or R1b1-L278, the second might correspond to older European hunter-gatherer lineages, or they might be associated with the expansion of Corded Ware cultures from the Balkans. Other non-R-M207 samples from central Corded Ware cultures include the two oldest ones, of IJ and G2a-P15 lineages, from Jagodno ca. 2800 BC[Gworys et al. 2013]; and one of haplogroup P-P295 in Esperstedt from ca. 2275 BC.
In the Baltic and the Forest zone, four samples are dated around 2500 BC: haplogroups R1a1a1b-Z645 and R1a1a1b1-Z283 in Kunila, and two samples of haplogroup R1a1a1b-Z645 in Ardu. These samples together with its previous presence in Usvyatyan culture (ca. 2500 BC) and in Naumovo and Sertaya II[Chekunova et al. 2014], and its continuity in later times suggest that R1a1a1b-Z645 lineages almost fully replaced the previous R1b-M343 lineages in the eastern Baltic around the time of the Corded Ware culture expansion.
The oldest R1a1a1-M417 lineages of Central Europe are found in early Corded Ware groups, while ancient DNA from Neolithic Linear Pottery (ca. 5500–3500 BC) and Globular Amphorae (ca. 3400-2800 BC) cultures have been found to correspond mainly to I2-M438 and G2a-P15 lineages, with no steppe-related ancestry in admixture analyses[Haak et al. 2015][Mathieson et al. 2017]. This raises the possibility of a wide and rapid spread of R1a1a1-M417, and especially R1a1a1b1-Z283 subclades in Europe[Underhill et al. 2015] associated with the expansion of the Corded Ware culture. The diversification of R1a1a1b2-Z93 in the Middle East and South Asia remains more obscure[Underhill et al. 2015].
However, in samples obtained from Estonia it was seen that admixture between Corded Ware culture farmers and Comb Ceramic culture hunter-gatherers may have been limited in males of R1a1a1-M417 lineages. Also, the presence of a genetic component associated with Caucasus hunter-gatherers – also present in Yamna migrants, Eastern hunter-gatherers, and individuals from the Estonian Comb Ceramic culture, means that the expansion of the Corded Ware culture cannot be seen as the sole means for the spread of this genetic component, at least in eastern Europe[Saag et al. 2017].
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