Middle East farmers
Hunters from the Pontic-Caspian steppe – as European Mesolithic hunter-gatherers in general – possessed no domesticated animals before the arrival of stockbreeding, spread with Neolithic farmers from Anatolia after about 6100 BC, probably through different colonisation routes through the Aegean, that involved diverse Neolithisation processes[Horejs et al. 2015].
While economic practices changed, some cultural traits like funerary practices did not accompany the ‘Neolithic package’ acquired in the Balkans by local fisher-hunter-gatherers. It seems that farmers and their domestic animals spread fast, in ca. 10 human generations, from sub-Mediterranean Macedonia to the northern limits of the temperate Balkan Peninsula and the adjacent Carpathian Plain, which may have put serious difficulties for the spread of cattle until selective pressure could drive genetically-driven adaptations to harsh environments[Ethier et al. 2017].
It has been supported with human ancestry studies that Middle East farmers arrived into central and western Europe with the Neolithic expansion[Brandt et al. 2013][Olalde et al. 2015][Szecsenyi-Nagy et al. 2017]. Ancient Middle Easterners show basal Eurasian ancestry with significantly less Neanderthal inheritance than East Asians, which suggests an affinity between Natufians and populations of north or sub-Saharan Africa. This is supported by the Y-DNA haplogroup E found in Levantine Neolithic populations[Lazaridis et al. 2016].
Together with WHG and EHG, samples from Neolithic Levant and those from the Neolithic of western Iran form the four streams of ancestry seen in the Middle East. There is continuity of Natufians (Levantine hunter-gatherers) with Levantine farmers, and Caucasus hunter-gatherers with farmers from the Zagros Mountains (east Iran).
While a population related to north-western Anatolian Neolithic farmers spread westward into Europe, farmers related to those of the Levant spread southward into north-east Africa[Lazaridis et al. 2016][Schuenemann et al. 2017].
Neolithic farmers arrived in Europe from a single Balkan population in two routes, one to the Linearbandkeramik (LBK) culture from central Europe, and another to the Impressa complex of Croatia and Epicardial Early Neolithic from Spain[Mathieson et al. 2017]. However, there are also traces of an outgroup related to the north-western Anatolian Neolithic population in the Peloponnese Neolithic, maybe linked to the pre-pottery Neolithic of Cyprus and the Levant. While farmers and hunter-gatherers lived in settlements in close proximity during the Neolithic, (in the Balkans, in western, central and northern Europe), there are signs of long periods with minimal admixture[Mathieson et al. 2017].
During the Middle Neolithic, a resurge of male-biased hunter-gatherer ancestry is seen in central Europe and Iberia, while persistent frontiers between hunter-gatherers and farmers are found in central and northern Europe, coincident with the loess belt of the northern European plain, to the north of which early farming techniques were probably not suitable. It is likely that new climates and environments led to the eventual breakdown of demic diffusion, and the spread of Neolithic traits by cultural diffusion[González-Fortes et al. 2017].
That resurgence of hunter-gatherer ancestry, with a ca. 4:1 WHG:EHG contribution, is found in the Balkan Neolithic in the territory of present-day Bulgaria, close to the Danube river. This suggests a heterogeneous landscape of farmer populations with different proportions of hunter-gatherer ancestry during the early Neolithic, probably due to pockets of hunter-gatherers surviving close to the coast and major rivers[Mathieson et al. 2017].
Image modified from Mathieson et al. (2017). «Structure and population change in European populations with farmer ancestry. A: each population is modeled as a mixture of Anatolia Neolithic, WHG, and EHG. Dashed lines show temporal relationships between populations from the same geographic areas with similar ancestries. B: Z-scores for the difference in hunter-gatherer ancestry on the autosomes compared to the X chromosome when populations are modeled as a mixture of Anatolia Neolithic and WHG. A positive score indicates that there is more hunter-gatherer ancestry on the autosomes and therefore the hunter-gatherer ancestry is male-biased». Original under a CC-BY-NC 4.0 International license.
The Pontic-Caspian steppe
Before the arrival of farmers to the western frontier of the Pontic-Caspian steppe, pottery was produced in the first half of the 7th millennium BC by hunter-gatherer groups first in the Volga steppes (with the earliest pottery found to date in the Elshanian culture). This culture was probably derived from the Eastern Asian tradition of the Late Pleistocene through Siberia and the Transurals[Piezonka 2015].
The first Neolithisation of the Lower Volga region, with the oldest pottery of ca. 6200 BC, can be attributed to the influence of the Kairshak culture in the northern Caspian region, where the first sites with the oldest pottery appeared ca. 6500 BC[Vybornov 2016]. From the north-western Caspian region pottery spread south- and westward into north Pontic societies ca. 6200-6000 BC[Zaitseva et al. 2009].
Sparsely decorated pottery dispersed north into the Forest Zone ca. 6000 BC or slightly earlier, from the upper Volga and Dvina-Lovat’ regions to the east (into the Dvina-Pechora region) and west (into the eastern Baltic), reaching the Upper Volga, Serteya, and Valday cultures, and later the Narva culture.
Contacts of north Pontic cultures with Criş settlers from the Starčevo–Kőrös–Criş culture about 5800 BC introduced domesticated cattle to the Bug-Dniester culture, but no signs of cultural assimilation has been found, with the later invasion of Linear Pottery sites ca. 5500-5200 BC respecting a similar cultural frontier, geographically coincident with the Dniester[Anthony 2007]. Hence the language of western Neolithic settlers – assumed to come from the Middle East, if language accompanied the spread of Neolithic technology – was probably not transferred to north Pontic herders.
A second expansion of eastern pottery reached the eastern Baltic region ca. 5500 BC, expanding from the Dnieper region to the north-west, generating the sparsely decorated Dubičiai pottery (later evolving into the Neman culture), and influencing the north European regions from the Narva to the Ertebølle cultures[Piezonka 2015].
From the Bug-Dniester culture domesticated cattle, sheep, and goats spread quickly from about 5200 BC east- and northward into Pontic-Caspian sites, reaching Khvalynsk and the Samara region about 5100 BC.
A third expansion of eastern pottery spread from the Volga-Kama region to the east ca. 5000 BC, connected to influences from beyond the Urals, showing a more elaborately decorated ware (with bands of pits and impressions made from comb stamps), spreading north and west in the Sperrings and Säräisniemi 1 cultures[Piezonka 2015].
A sample of R1b1a1a-P297 reported as possibly an intermediate stage of its formation (positive and negative markers in the M478 node) was found in a Mesolithic hunter-gatherer at Lebyanzhinka in the Samara region, dated ca. 5600 BC[Mathieson et al. 2015]. Later samples from the same region show continuity of R1b1a1a2-M269 lineages, which seem to have expanded from east to west in the Pontic-Caspian steppes.
In the north Pontic steppe – apart from the earlier R1a-M420 and R1b-M343 samples from Mesolithic Vasylivka (see above) – there are four samples (dated ca. 6500-4000 BC) of the Mariupol culture from Volniensky, and one sample of the Azov-Dnieper culture from Vovnihi (ca. 5400 BC), of I-M170 (and one IJ) lineage.
Samples attributed to the early Sredni Stog culture in Deriivka (dated ca. 5500-4800 BC) include nine of R1b-M343 lineage, probably from an extinct branch of R1b1a-L754 (xR1b1a1a-P297, xR1b1a1a2-M269); one of R1a-M420 lineage; and four samples of haplogroup I-M170, probably I2a2a1b-L701[Mathieson et al. 2017].
This diversity of lineages points to a mix in the different groups that emerged during the Mesolithic and early Neolithic periods, before the mass expansions that occurred later.
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