Alternative models of Indo-European expansion
This model uses mainly SNP investigation to support the general view of Archaeologists whereby Yamna migrants did not directly form the Corded Ware culture[Anthony 2007][Harrison and Heyd 2007][Heyd 2007][Heyd 2012][Kristiansen et al. 2017][Heyd 2017], correcting thus assumptions based solely on recent genetic research.
Communities formed mainly by R1b1a1a2a-L23 and R1a1a1b-Z645 lineages, which clearly expanded during the Chalcolithic – associated to Yamna and Corded Ware cultures, respectively – are assumed to have evolved differently in eastern Europe, albeit in close contact (probably in a neighbouring region) in light of the common ancestry they share.
Other models are compatible with the state of the art of genetic investigation, though. Listed here are possibilities compatible with the current knowledge, in decreasing order of probability:
Yamna clans of R1a1a1b-Z645 lineages existed and migrated to the north, but have not been sampled yet. This seems to be the preferred model in recent genetic papers, which support a direct Yamna > Corded Ware migration[Haak et al. 2015][Allentoft et al. 2015][Lazaridis et al. 2016].
This framework seems compatible with Kristiansen’s traditional model of Corded Ware culture development[Kristiansen 1989][Kristiansen et al. 2017], and also partially with that of Gimbutas[Gimbutas 1965][Gimbutas 1977], involving a gradual cultural (and population) contribution of Indo-Europeans from the steppe to east-central Europe. Centuries of contact between the Indo-Europeans from the steppe and the Baden and Globular Amphora cultures should have formed the third Corded Ware horizon that expanded after ca. 3000 BC.
This seems in part supported by steppe ancestry found in individuals from the Balkans at Varna I (ca. 4630 BC), Smyadovo (ca. 4500 BC), and outliers from Ukraine Eneolithic (ca. 3500 BC) and Trypillian culture (ca. 3325 BC). All of them clearly show such steppe ancestry before the main Chalcolithic expansion[Mathieson et al. 2017][Haak et al. 2015].
However, it is in contrast with the lack of steppe ancestry in sampled distant individuals from Baden and Globular Amphora cultures[Mathieson et al. 2017]. Also, from a linguistic point of view, this long-lasting framework does not allow for the adoption of a Late Indo-European language by the Corded Ware culture, and would still leave the most obvious Late Indo-European-speaking expansion to the spread of Yamna migrants.
On the other hand, the model presented in this paper, which supports a Corded Ware Substratum Hypothesis, would be compatible with eastern Yamna clans of R1a1a1b-Z645 lineages expanding a language ancestral to Slavic and Indo-Iranian with the Corded Ware culture, and therefore the substrate language would correspond to the original east-central region where the culture originated in the Late Neolithic. In that case, Kortlandt’s late Sətem Indo-Slavonic dialect[Kortlandt 2016b] could also be supported, but the common unconnected substratum of Balto-Slavic and Germanic[Kortlandt 2016a] would probably need further assumptions, like the Agricultural Substrate Hypothesis[Kroonen 2012][Kristiansen et al. 2017][Iversen and Kroonen 2017].
Problems with this model include:
- It assumes the existence of clans with a majority of R1a1a1b-Z645 lineages coexisting with clans with a majority of R1b1a1a2a1-L51 subclades, either in the east or in the west – or, exceptionally, in both, since they are different dialectal areas. Both of them would have expanded during the Chalcolithic: R1a1a1b-Z645 clans to the north (possibly via the Prut), to form the third Corded Ware horizon, and R1b1a1a2a1-L51 and R1b1a1a2a2-Z2103 clans to the east into the Afanasevo culture, and west to form Yamna settlements of the Balkans.
- It would need a framework for a sudden and direct contribution of Yamna to the creation of the Corded Ware culture, which currently does not exist.
- The coexistence of such differentiated communities is in contrast with the natural evolution of male-dominated steppe societies of R1b1a1a2-M269 and R1b1a1a2a-L23 lineages, which underwent at least two important expansions, and must have therefore undergone a decrease in Y-DNA variability since the Mesolithic.
- It is also in contrast with the lack of R1a1a1b-Z645 lineages in Yamna, and in western and south-eastern European cultures related to the expansion of Indo-European languages. A potential explanation for this lack of R1a1a1b-Z645 would be a sample selection bias, whereby western Yamna migrants of R1a1a1b-Z645 subclades and central-east European Corded Ware samples of R1b1a1a2a-L23 subclades have not yet been sampled.
- A further assumption of this model is that Corded Ware contributed to the creation of the Bell Beaker culture[Haak et al. 2015][Kristiansen et al. 2017], which is in contrast with mainstream archaeological models, and is based mainly on the influence of the Corded Ware outlier of Erperstedt in assessing steppe ancestry of the Corded Ware culture. The closer position of Bell Beaker samples from the Balkans to Yamna samples – closer than any other sample of the Corded Ware in PCA[Olalde et al. 2017] –, as well as their connection to Vučedol and western Yamna ancestry, makes such a direct connection unlikely.
Founder effect / resurgence of R1a lineages or cultural diffusion
West Yamna was mainly composed of clans of R1b1a1a2a-L23 subclades, as supported in this model, but the admixture seen in Corded Ware samples from central and north-eastern Europe comes precisely from original Yamna migrants of R1b1a1a2a-L23 subclades that have not been sampled. These first generations of northern migrants would have travelled west and north-west, maybe involving those known to have settled north up the Prut River, possibly beginning as early as 3100 BC, but probably from ca. 3000 BC.
On the other hand, Anthony’s model would still need a majority of the Usatovo population mainly composed of R1a1a1b-Z645 lineages, and with a previous ancestry closest to the east Yamna population, as well as a resurge of R1a1a1b-Z645 lineages within few generations, so that admixture and Yamna lineages could go unnoticed in the Corded Ware population sampled. To sample individuals from only a few generations is difficult, apart from the indemonstrable proposal (advanced by Anthony) that the general Usatovo population would have adopted the language of the elites (in this case Pre-Germanic) by way of cultural diffusion. It also leaves open the same possibility for multiple (as unlikely and as indemonstrable) cultural diffusions among Corded Ware groups, e.g. of Proto-Balto-Slavic to the Middle Dnieper culture.
R1b-L51 from the west and cultural diffusion
West Yamna clans were mainly composed of clans of R1b1a1a2a2-Z2103 lineages, which expanded with Yamna migrants to the west. Subclade R1b1a1a2a1-L51 (and especially R1b1a1a2a1-L151 subclades) may have split after the Neolithic expansion associated with Middle Indo-European, and developed a society in central – south-east Europe.
It would have later adopted the ‘Yamna package’, developing the East Bell Beaker group that later expanded further to the west. That could be supported by the finding of a R1b1a1a2a2-Z2103 in Vučedol, and the lack of R1b1a1a2a1-L51 subclades in Yamna. This model could be linked with the proposed origin of Proto-Indo-European in western or central Europe[Cunliffe and Koch 2012].
This is in contrast with estimated dates of haplogroup formation, and the known history from ancient samples. The evolution of R1b1a1a2-M269 and R1b1a1a2a-L23 lineages within the steppe since the Mesolithic, the quite late TMRCA for R1b1a1a2a-L23, and the finding of an R1b1a1a2a-L23(xZ2013) subclade in eastern Yamna probably points to an internal evolution of these subclades. This and the expansion of R1b1a1a2a1-L151 subclades (but not R1b1a1a2a1-L51) with the Bell Beaker culture, probably evolved from Yamna migrants in the westernmost region, makes the eventual appearance of R1b1a1a2a1-L51 subclades in west Yamna quite likely.
This model would also allow for a Yamna region hiding unsampled clans of R1a1a1b-Z645 lineages, as in the Kurgan people model.
An alternative hypothesis is that the Proto-Indo-European homeland was in the Caucasus or Iran, and expanded through Anatolia, which can be linked with the most recent proposals of the Anatolian hypothesis[Renfrew 2003]. The current genetic models are not, however, compatible with the Armenian homeland hypothesis[Gamkrelidze and Ivanov 1995], which suggest a more recent expansion of Late Indo-European from the Armenian highland. A westward movement associated with the CHG ancestry may then have contributed to the dispersal of Anatolian languages, seen in the contribution of CHG to Anatolian peoples during different Neolithic and Chalcolithic waves. The northward mixture of CHG with EHG ancestry in the steppe may signal the formation of Indo-European-speaking steppe population associated with the Chalcolithic expansion of Late Indo-European[Mathieson et al. 2017][Lazaridis et al. 2017].
Haplogroup analysis reveals a prevalence of J-M304 lineages associated with CHG expansion into Anatolia, probably related to movements within the Fertile Crescent, which is not seen in the Yamna population. This suggests a different type of spread for this component in the steppe.
No archaeological models show such a strong wave of migrants from the south into the steppe, but continuous contacts of steppe cultures with Transcaucasian cultures since the Mesolithic. It is therefore more likely that the CHG contribution came from a long-term inter-regional gene flow that began early, probably coinciding with Neolithic population movements within the steppe.
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