This paper presents the first comprehensive pan-Iberian overview of one of the major episodes of cultural change in later prehistoric Iberia, the Copper to Bronze Age transition (c. 2400–1900 BC), and assesses its relationship to the 4.2 ky BP climatic event. It synthesizes available cultural, demographic and palaeoenvironmental evidence by region between 3300 and 1500 BC. Important variation can be discerned through this comparison. The demographic signatures of some regions, such as the Meseta and the southwest, diminished in the Early Bronze Age, while other regions, such as the southeast, display clear growth in human activities; the Atlantic areas in northern Iberia barely experienced any changes. This paper opens the door to climatic fluctuations and inter-regional demic movements within the Peninsula as plausible contributing drivers of particular historical dynamics.
Interesting excerpts summarizing key trends in the different regions:
Between 2200 and 1900 BC, the northernmost regions (i.e. Galicia, the Cantabrian strip and the northeastern sector to the north of the Ebro valley) underwent relatively minor changes in the realms of settlement and burial practices. (…) In addition, some Atlantic areas show a marked and statistically significant fall in human activity c. 2200 BC, with a subsequent recovery c. 1600 BC, and such observations are matched by paleoenvironmental proxies and a lack of known EBA sites
The overall impression from the Meseta is one of sharp disruption in cultural practices; these include both settlement and burial patterns, abrupt shifts in local climate conditions, and striking differences in human pressure on vegetation. However, there was also clear intra-regional variability, with remarkable internal particularities and differential tempos between the western and eastern sectors. In terms of material culture, discontinuity with the Copper Age is the main trend in the western Duero and the Tagus valleys, yet EBA communities to the north of the Central System adopted far more distinctive and therefore traceable site types (hilltops) and material repertoire. This shift was even stronger in the case of the Motillas culture at La Mancha, whose pathway seems closely tied to the Argaric area.
Intra-regional variability is also apparent within the northeast (…) In the second millennium BC, material culture changed, long-distance exchange intensified and anthropogenic pressure increased, despite continuity in diverse realms of social practice.
The pattern in the southwest was one of marked discontinuity with two key features: a) it follows the general decreasing trends manifest across Atlantic Iberia; and b) its temporality was clearly different from the rest of the Peninsula and apparently unrelated to the 4.2 ky BP event. Thus, a highly conspicuous and rich variety of cultural expressions in the Chalcolithic, with an early and marked peak in human activity during the Beaker phase c. 2500 BC, gave way to a sudden cultural collapse prior to the onset of the EBA
The southeast exhibits one of the most remarkable cultural shifts in Western Europe. (…) The radical transformation in Chalcolithic materiality and ways of life could be regarded as a kind of societal collapse. The Argaric, a highly hierarchical and integrated regional polity, is the clearest example of a new scenario that emerged after the 4.2 ky BP event, yet the contributing role of environmental change and immigration from other regions remains to be fully explored.
Since R1b-DF27 lineages are widely distributed in modern western Europe, it is only logical that the recent find of its first ancient sample in Iberia has sparked the interest for Chalcolithic and Early Bronze Age Iberian cultures.
There is not much literature in English about Iberian prehistory, especially on the evolution of Bell Beaker culture. Also, most papers in Spanish on this cultural phenomenon – in my humble opinion, as a non-archaeologist – seem to be written from a merely descriptive archaeological point of view, many of them still sharing the radiocarbon-based assessment of origin and distribution of materials, instead of more complex anthropological models of cultural change and potential migrations.
Nevertheless, changes and influences in Iberian cultures are obvious regardless of the view taken on population movements (which are becoming quite clear now), and this paper seems to me a thorough review, very interesting for international researchers when interpreting ancient DNA from Iberia.
Featured image, modified from the paper: “The Bell Beaker culture in the northern Meseta: an artistic recreation of funerary ritual at Fuente Olmedo (Valladolid). Source: Garrido-Pena et al. 2011, Fig. 7.7”.
Population genomic studies of ancient human remains have shown how modern-day European population structure has been shaped by a number of prehistoric migrations. The Neolithization of Europe has been associated with large-scale migrations from Anatolia, which was followed by migrations of herders from the Pontic steppe at the onset of the Bronze Age. Southwestern Europe was one of the last parts of the continent reached by these migrations, and modern-day populations from this region show intriguing similarities to the initial Neolithic migrants. Partly due to climatic conditions that are unfavorable for DNA preservation, regional studies on the Mediterranean remain challenging. Here, we present genome-wide sequence data from 13 individuals combined with stable isotope analysis from the north and south of Iberia covering a four-millennial temporal transect (7,500–3,500 BP). Early Iberian farmers and Early Central European farmers exhibit significant genetic differences, suggesting two independent fronts of the Neolithic expansion. The first Neolithic migrants that arrived in Iberia had low levels of genetic diversity, potentially reflecting a small number of individuals; this diversity gradually increased over time from mixing with local hunter-gatherers and potential population expansion. The impact of post-Neolithic migrations on Iberia was much smaller than for the rest of the continent, showing little external influence from the Neolithic to the Bronze Age. Paleodietary reconstruction shows that these populations have a remarkable degree of dietary homogeneity across space and time, suggesting a strong reliance on terrestrial food resources despite changing culture and genetic make-up.
We present a comprehensive biomolecular dataset spanning four millennia of prehistory across the whole Iberian Peninsula. Our results highlight the power of archaeogenomic studies focusing on specific regions and covering a temporal transect. The 4,000 y of prehistory in Iberia were shaped by major chronological changes but with little geographic substructure within the Peninsula. The subtle but clear genetic differences between early Neolithic Iberian farmers and early Neolithic central European farmers point toward two independent migrations, potentially originating from two slightly different source populations. These populations followed different routes, one along the Mediterranean coast, giving rise to early Neolithic Iberian farmers, and one via mainland Europe forming early Neolithic central European farmers. This directly links all Neolithic Iberians with the first migrants that arrived with the initial Mediterranean Neolithic wave of expansion. These Iberians mixed with local hunter-gatherers (but maintained farming/pastoral subsistence strategies, i.e., diet), leading to a recovery from the loss of genetic diversity emerging from the initial migration founder bottleneck. Only after the spread of Bell Beaker pottery did steppe-related ancestry arrive in Iberia, where it had smaller contributions to the population compared with the impact that it had in central Europe. This implies that the two prehistoric migrations causing major population turnovers in central Europe had differential effects at the southwestern edge of their distribution: The Neolithic migrations caused substantial changes in the Iberian gene pool (the introduction of agriculture by farmers) (6, 9, 11, 13, 24), whereas the impact of Bronze Age migrations (Yamnaya) was significantly smaller in Iberia than in north-central Europe (24). The post-Neolithic prehistory of Iberia is generally characterized by interactions between residents rather than by migrations from other parts of Europe, resulting in relative genetic continuity, while most other regions were subject to major genetic turnovers after the Neolithic (4, 6, 7, 9, 25, 48). Although Iberian populations represent the furthest wave of Neolithic expansion in the westernmost Mediterranean, the subsequent populations maintain a surprisingly high genetic legacy of the original pioneer farming migrants from the east compared with their central European counterparts. This counterintuitive result emphasizes the importance of in-depth diachronic studies in all parts of the continent.
The roles of migration, admixture and acculturation in the European transition to farming have been debated for over 100 years. Genome-wide ancient DNA studies indicate predominantly Anatolian ancestry for continental Neolithic farmers, but also variable admixture with local Mesolithic hunter-gatherers. Neolithic cultures first appear in Britain c. 6000 years ago (kBP), a millennium after they appear in adjacent areas of northwestern continental Europe. However, the pattern and process of the British Neolithic transition remains unclear. We assembled genome-wide data from six Mesolithic and 67 Neolithic individuals found in Britain, dating from 10.5-4.5 kBP, a dataset that includes 22 newly reported individuals and the first genomic data from British Mesolithic hunter-gatherers. Our analyses reveals persistent genetic affinities between Mesolithic British and Western European hunter-gatherers over a period spanning Britain’s separation from continental Europe. We find overwhelming support for agriculture being introduced by incoming continental farmers, with small and geographically structured levels of additional hunter-gatherer introgression. We find genetic affinity between British and Iberian Neolithic populations indicating that British Neolithic people derived much of their ancestry from Anatolian farmers who originally followed the Mediterranean route of dispersal and likely entered Britain from northwestern mainland Europe.
Quite interesting are the early samples from Poland, of R1b1a1a2a2-Z2103 and R1b1a1a2a1a-L151 lineages – , which may point (different to the more homogeneous L151 distribution in Western Europe) to a mix in both original (east-west) Yamna groups. This could tentatively be used to explain the Graeco-Aryan influence that some linguists see in Balto-Slavic (or its superstrate).
That link would then be quite early, to account for an influence during the Yamna settlements in Hungary, before its expansion as East Bell Beakers, but we haven’t seen a clearly differentiated subgroup (yet) in Archaeology, Anthropology, or Genomics within the Hungarian Yamna/East Bell Beaker community, so I am not convinced. It could be just that different scattered subclades mixed with the general L151 population pop up (following old Yamna lineages, or having being added along the way), as expected in an expansion over such a great territory – as if some scattered samples of R1a, I1, I2, J, etc. were found.
We need more early samples from south-eastern Europe and the steppe during the Chalcolithic to ascertain the composition and migration paths of the different Yamna settlers.
I mentioned in the Indo-European demic diffusion model the need to assess absolute and relative population growth – as well as other demographic changes – to interpret genomic data from the different European regions studied.
The neolithization of Northern and Northwestern Europe (probably with new forms of slash-and-burn agriculture; Feeser et al. 2012; Schier 2009) was also one of the causes for the population increase observed.
The introduction of the plough and developing technologies (e.g. the introduction of the wheel) (cf. Mischka 2011) might also be causes of rising population figures from ca. 3500–3000 BC.
The establishment of subcontinental value systems, such as the Corded Ware and Bell-Beaker phenomena (Czebreszuk/Szmyt 2003; Furholt 2004), in contrast to regional identities, might have triggered different reactions in different areas, leading to fluctuating population levels.
The introduction of Bronze Age ideologies, including bronze as a technology, triggered the spread of Neolithic and Bronze Age societies to vast areas of Europe (e.g. Earle/Kristiansen 2010).A major population increase is observed in both the areas already settled as well as in new areas of interest.
In our population estimations for Central Europe and Scandinavia, population increases are associated with the periods from 5500–5000 BCE (LBK) and 3500–3000 BCE (middle and late Funnelbeaker Culture), but not for the period from 2500–2000 BCE (Bell Beaker). Consequently, this would possibly indicate forms of immigration for the first two periods and a form of interregional networking (e.g. through marriage) for the latter. But as also for the first cases on the supra-regional level (which our enquiries investigated), no other area with a significant population decrease could be observed, therefore “proof” for larger population displacement is not given. For such inquiries, studies on a more regional level are probably necessary. Nevertheless, for the Bell Beaker period I would like to exclude the possibility of large population influxes at least to Central Europe and South Scandinavia as the population values show no indication of such an event (cf. Fig. 10). In consequence, supra-regional networks or population-exchanges between smaller regions might be responsible for the isotope values.
One could interpret from the graphics, including known anthropological and genomic data, that:
The Yamna expansion into South-East Europe must have included some population replacement, i.e. influx into progressively deserted areas (such as that of the Cucuteni-Trypillia culture), since it did not leave traces of population growth.
The impact of the expansion of East Bell Beakers from ca. 2500 BC is clear in South-East Europe, and especially in Western Europe – taking into account the whole population growth in Europe. In Central Europe and Scandinavia the overall impact of BB migration was more limited, which suggests some degree of population replacement.
Also important to interpret genomic data are the actual economic and social differences in the different periods and cultures – usually growing after the introduction of farming. A good example is the scarce data from Khvalynsk, where the sample of haplogroup R1b (most likely of subclade M269) shows – apart from a closer position in PCA to Yamna – a a high-status burial, similar to high-status individuals buried under kurgans in later Yamna graves. This man was therefore probably a founder of an elite group of patrilineally-related families, which dominated in the following Yamna culture, which explains the clear expansion of this haplogroup’s subclades from this region.
Other interesting papers on European demographics by Johannes Müller include:
Featured image, from the main article: “The distribution of agrarian regions in Europe and the Near East in relation to the supra-regions as defined in this study: Near East (NE) about 2.400.000 km2; South East Europe (SEE) about 1.087500 km2; Central Europe and South Scandinavia (CE/SSc) about 1.613.000 km2. Europe includes 10.050.000 km2 (without Iceland)”.
Important excerpts for the Indo-European question (emphasis mine):
Mesolithic to Neolithic
In the archaeological understanding, the transition from Mesolithic to Neolithic in the Eastern Baltic region does not coincide with a large-scale population turnover and a stark shift in economy as seen in Central and Southern Europe. Rather, it is signified by a change in networks of contacts and the use of pottery, among other material, cultural and economic changes. Our results suggest continued admixture between groups in the south of the Eastern Baltic region, who are more closely related to WHG, and northern or eastern groups, more closely related to EHG. Neolithic social networks from the Eastern Baltic to the River Volga could also explain similarities of the hunter-gatherer pottery styles, although morphologically analogous ceramics might also have developed independently due to similar functionality. The genetic evidence for a change in networks and possibly even a large-scale population movement is most pronounced in the Middle Neolithic in individuals attributed to the CCC. The distribution of this culture overlaps in the north with the Narva culture and extends further north to Finland and Karelia. Its spread in the Eastern Baltic is linked with a significant change in imported raw materials, artefacts, and the appearance of village-like settlements15.
Neolithic to Chalcolithic
We see a further population movement into the regions surrounding the Baltic Sea with the CWC in the Late Neolithic that was accompanied by the first evidence of extensive animal husbandry in the Eastern Baltic. The presence of ancestry from the Pontic-Caspian Steppe among Baltic CWC individuals without the genetic component from north-western Anatolian Neolithic farmers must be due to a direct migration of steppe pastoralists that did not pick up this ancestry in Central Europe. It suggests import of the new economy by an incoming steppe-like population independent of the agricultural societies that were already established to the south and west of the Baltic Sea. The presence of direct contacts to the steppe could lend support to a linguistic model that sees an early branching of Balto-Slavic from a Proto-Indo-European language, for which the west Eurasian steppe was proposed as a homeland. However, as farmer ancestry is found in later Eastern Baltic individuals, it is likely that considerable individual mobility and a network of contact throughout the range of the CWC facilitated its spread eastward, possibly through exogamous marriage practices. Conversely, the appearance of mitochondrial haplogroup U4 in the Central European Late Neolithic after millennia of absence could indicate female gene-flow from the Eastern Baltic, where this haplogroup was present at high frequency.
There obviously was exogamy – which may in fact justify the findings in PCA close to Yamna (like the Zvejnieki sample), although researchers obviate that.
Also, as expected, no R1b-M269 in the Baltic (during the Corded Ware period), most are R1a with the majority showing subclade R1a-Z645 (and others poor SNP coverage), which support the reduction in haplogroup diversity to this very subclade during the expansion of Corded Ware peoples, as I predicted it would happen.
Local foraging societies were, however, not completely replaced and contributed a substantial proportion to the ancestry of Eastern Baltic individuals of the latest LN and Bronze Age. This ‘resurgence’ of hunter-gatherer ancestry in the local population through admixture between foraging and farming groups recalls the same phenomenon observed in the European Middle Neolithic and is responsible for the unique genetic signature of modern-day Eastern Baltic populations.
We suggest that the Siberian and East Asian related ancestry in Estonia, and Y-haplogroup N in north-eastern Europe, where it is widespread today, arrived there after the Bronze Age, ca. 500 calBCE, as we detect neither in our Bronze Age samples from Lithuania and Latvia. As Uralic speaking populations of the Volga-Ural region show high frequencies of haplogroup N, a connection was proposed with the spread of Uralic language speakers from the east that contributed to the male gene pool of Eastern Baltic populations and left linguistic descendants in the Finno-Ugric languages Finnish and Estonian. A potential future direction of research is the identification of the proximate population that contributed to the arrival of this eastern ancestry into Northern Europe.
It is funny to see how people keep trying to identify R1a with ‘Yamnaya’, now ‘steppe’, but always Indo-European (an ethnolinguistic term, mind you) supposedly because of the ‘Yamnaya’ (now ‘steppe’) admixture, but the only ‘mark’ of Uralic languages for the same researchers in the same paper using this very concept is nevertheless, paradoxically, haplogroup N, with an assumption explicitly based on prevalence in modern populations…
This admixture vs. haplogroup questionfor language and culture identification in genetic papers is really gettting messed up with new data, now in a contortionist-like way…
Images and text: Content of the paper is licensed under CC-by 4.0.
Recent genetic studies have claimed to reveal a massive migration of the bearers of the Yamnaya culture (Pit-grave culture) to the Central and Northern Europe. This migration has supposedly lead to the formation of the Corded Ware cultures and thereby to the dispersal of Indo-European languages in Europe. The article is a summary presentation of available archaeological, linguistic, genetic and cultural data that demonstrates many discrepancies in the suggested scenario for the transformations caused by the Yamnaya “invasion” some 5000 years ago.
Both teams [Reich/Anthony, and Willerslev/Kristiansen] interpreted this resemblance in the same way: as evidence of mass migration of the Yamnaya culture from the steppes into the Central and Northern Europe, resulting in the formation of the Corded Ware cultures, and these are universally recognised as Indo-European. Since earlier in this part of Europe existed a different pool of genomes, geneticists presumed that the Yamnaya migration alone had brought the Indo-European languages into Europe. It is difficult to say to what extent the pre-convictions of the involved archaeologists influenced these conclusions, or whether the results of the genetic studies attracted archaeologists with such beliefs.
Mismatch of cultural manifestations
First, we might question the idea of the Yamnaya culture as a unity rather than a loose conglomerate of cultures. Merpert (1974) divided it into nine local groups but did not recognise them as separate cultures. However, in 1975 I suggested that Nerushay (Budzhak) monuments should be recognised as a distinct culture (Klejn 1975), although still as a part of the same broader steppe community.
This was accepted by other specialists (Ivanova 2012; 2013; 2014). Generally, in the western branch of this community, a mixture of the eastern rites of interment with local, Balkan ceramics can be observed. It should be noted that hitherto all genetic samples were taken from eastern material (in the vicinity of Samara in the Volga basin and Kalmykia), while the central thesis concerns the intrusion of the western branch of this community (Budzhak culture) into Europe.
Simultaneity of cultures
The Yamnaya culture (Chernykh & Orlovskaya 2004a; Heyd 2011; Frȋnculeasa et al. 2015) appears not to be the predecessor of the Corded Ware cultures but is contemporary with them. The Corded Ware cultures appeared also around the turn between the fourth and third millennium BC (Stöckli 2001; Furholt 2003). Their derivation from the Yamnaya seems, therefore, to be less probable. This is evidenced by the fact that the corded beakers or amphorae found in the Budzhak culture are not the prototypes of the corded beakers or amphorae found in more northern territories, but seem instead to be an outcome of contemporaneous contacts (Ivanova 2014; Klejn 2017c).
Discrepancies across the haplogroups
Even more remarkable is the variation in the distribution of types of Y chromosome. In the Yamnaya population, R1b is not just a single occurrence (there are about seven known occurrences) while in the Corded Ware population a different clade of R1b is found and R1a is predominant (several instances). Thus the postulate of unbroken succession finds no support!
In the tables presented in the article by Reichs’ team (Haak et al. 2015) the genetic pool connecting the Yamnaya culture with the Corded Ware people is shown to be more intense in Northern Europe (Norway and Sweden) and decreases gradually from the North to the South (Fig. 6). It is weakest around the Danube, in Hungary, i. e. areas neighbouring the western branch of the Yamnaya culture! This is the reverse image to what the proposed hypothesis by the geneticists would lead us to expect. It is true that this gradient is traced back from the contemporary materials, but it was already present during the Bronze Age (Klejn 2015a).
The author also uses questionable interpretations from selected articles to advance his (as of today) untenable positions regarding a Mesolithic origin of the reconstructible Proto-Indo-European language.
1. Glottochronology, for a PIE origin:
If based on the data of glottochronology (taking into account all disputes) the period of initial dispersal is to be dated to the 7th-5th millennium BC.
The currently available dataset does not contradict the hypothesis that R-GG400 marks a link between the East European steppe dwellers and West Asians, though the route and even direction of this migration is disputable. It does, however, demonstrate that present-day West European R1b chromosomes do not originate from the Yamnaya populations analyzed in (Haak et al. 2015; Mathieson et al. 2015) and raises the question of their origin. A Bronze Age origin is more likely than a Neolithic one (Balaresque et al. 2010), but further ancient DNA studies may be necessary to identify this source.
This is usual with amateur geneticists (those who don’t publish, and are therefore not subjected to criticism): if anyone is wrong (whether in Archaeology or Genetics), then they are wrong in everything else. It seems to me that Klejn’s theses against recent genetic results rest on the same assumption: The Yamna -> Corded Ware migration model is wrong, ergo the Yamna homeland model is wrong.
I guess this same fallacy is what a lot of angered geneticists (whether professional or amateurs) are going to use to dismiss Klejn’s criticism, trying to focus on what he clearly does not grasp – about genomic data of Yamna peoples and their expansion – to disregard his doubts on genetic interpretations entirely.
I have warned many times about how simplistic interpretations of genetic data would cause a general mistrust in the field, and that archaeologists won’t take the discipline seriously, no matter how many articles get published in famous research tabloids like Nature or Science…
Those who dismiss this warning lightly seem to forget the fate of other recent “scientific breakthroughs” which were initially so promising that Humanities appeared to matter no more, like glottochronology for Linguistics and, to some extent, that of radiocarbon analysis for Archaeology. EDIT: see here a recent example of discusion on discrepancies between archaeological and 14C-based chronologies, whereby ‘scientific data’ obviously needs archaeological context for a meaningful interpretation
Featured image: The direction of the supposed migration of the bearers of the Yamnaya culture into the area of the Corded Ware cultures. After Haak et al. 2015.
NOTE: I obviously don’t agree with Klejn’s main model: he criticises the Proto-Indo-European steppe homeland, and more specifically the expansion of Yamna peoples with R1b-L23 subclades, which I support. But, probably because of his “pre-convictions” (as he puts it when describing proponents of the steppe hypotheses) about the Proto-Indo-European homeland in Northern Europe during the Mesolithic, he was one of the first renown archaeologists to criticise the obvious inconsistencies in the genetic model of migrations based exclusively on the “Yamnaya ancestral component” concept, and to provoke the necessary reaction from (until then) overconfident geneticists, and he deserves credit for that.
His recent studies include important sites (for Archaeology and recently also for Genomics) such us Dereivka and Alexandria, part of the North Pontic steppe and southern steppe-forest zone, on the Left Bank of the Dnieper river. According to him, many of these sites seem to form part of a common and distinct cultural group.
The author discusses the issues of chronology of the known burial grounds. In this article, first of all, the location of a series of burials at Ihren 8 cemetery is revised and the earlier proposed point of view of the author himself is refined. An important moment for that was a revision of a paired bi-ritual burial 7-8 from the excavations in 1932 with the Trypillian painted cup of the second half of the Trypillia B/2. The author presents the arguments for the assumption that the two burials were made not at the same time. As a whole, singled out are the Early Chalcolithic burials with the peculiar for them position on a back with bended knees, accompanied by flint products, first of all tools made on long blades. The second later group is represented by two supine burials which date is determined by a Trypillian cup. Concerning Oleksandriia burial ground, the author confirms his earlier expressed position on the Early Chalcolithic age of the burials with long flint blades, presenting additional arguments, one of which is a publication of a new radiocarbon date for one of the burials. Based on the author’s terminology, graves of the both burial grounds are considered within the borders of the so called Skelianska culture existence, while in Ihren burial ground several burials could be made in the period of so called «hiatus» when there were the Stohivska group sites in the Dnipro River region.
A new burial complex is publishing by authors. This burial complex finds analogies among the Early Eneolithic burials of the Siversky Donets basin according to the rite and inventory (long flint blade). In addition, a set of specific flint products (long blades, triangular «spear heads» and flat adzes) finds analogies at the Aleksandriia settlement, where Skelia-type ceramics are represented. Therefore, there is a reason to combine in the same cultural and chronological context the relevant materials of the Aleksandriia settlement and the Early Eneolithic burials, and consider their as a part of the phenomenon that one of the authors conventionally calls Skelia culture.
It remains to bee seen how this new data is interpreted with more complex anthropological models, of potential cultural-historical groups that might have shaped posterior migrations.
I already wrote about the concept of outlier in Human Ancestry, so I am not going to repeat myself. This is just an update of “outliers” in recent studies, and their potential origins (here I will repeat some of the examples):
Early Khvalynsk: the three samples from the Samara region have quite different positions in PCA, from nearest to EHG (of Y-DNA haplogroup R1a) to nearest to ANE ancestry (of Y-DNA haplogroup Q). This could represent the initial consequences of the second wave of ANE ancestry – as found later in Yamna samples from a neighbouring region -, possibly brought then by Eurasian migrants related to haplogroup Q.
With only 3 samples, this is obviously just a tentative explanation of the finds. The samples can only be reasonably said to show an unstable time for the region in terms of admixture (i.e. probably migration), judging by the data on PCA.
Ukraine Eneolithic samples offer a curious example of how the concept of outlier can change radically: from the third version (May 30th) of the preprint paper of Mathieson et al. (2017), when the Ukraine Eneolithic sample with steppe ancestry (and clustering with central European samples) was the ‘outlier’, to the fourth version (September 19th), when two samples with steppe ancestry clustering close to Corded Ware samples were now the ‘normal’ ones (i.e. those representing Ukraine Eneolithic population), and the outlier was the one clustering closely with Ukraine Mesolithic samples…
This is one of the funny consequences of the wrong interpretation of the ‘yamnaya component’, that made geneticists believe at first that, out of two samples (!), the ‘outlier’ was the one with ‘yamnaya’ ancestry, because this component would have been brought by an eastern immigrant from early Khvalynsk…
West Yamna (to insist on the same question, the ‘yamnaya’ component): we have only four western Yamna samples, two of them showing Anatolian Neolithic ancestry (one of them, from Ukraine, with a strong ‘southern’ drift). On the other hand, Corded Ware migrants do not show this. So we could infer that their migrations were not coetaneous: whereas peoples of Corded Ware culture expanded ca. 3300 BC to the north – in the natural corridor to the Baltic that has been proposed for this culture in Archaeology for decades (and that is well represented by Ukraine Eneolithic samples) -, peoples of Yamna culture expanded to the west, replacing the Ukraine Eneolithic population (i.e. probably those of ‘Proto-Corded Ware culture’), and eventually mixing with Balkan populations of Anatolian Neolithic ancestry.
Potapovka, Andronovo, and Srubna: while Potapovka clusters closely to the steppe, and Andronovo (like Sintashta) clusters closely to Corded Ware (i.e. Ukraine Neolithic / Central-East European), both have certain ‘outliers’ in PCA: the former has one individual clustering closely to Corded Ware, and the latter to the steppe. Both ‘outliers’ fit well with the interpretation of the recent mixture of Corded Ware peoples with steppe populations, and they offer a different image for the evolution of populations of Potapovka and Sintashta-Petrovka, potentially influencing their language. The position of Srubna samples, nearer to Sintashta and Andronovo (but occupying the same territory as the previous Potapovka) offers the image of a late westward conquest from Corded Ware-related populations.
Iron Age Bulgaria: a sample of haplogroup R1a-z93, with more ‘yamnaya’ ancestry than any other previous sample from the Balkans. For some, it might mean continuity from an older time. However – as with the Corded Ware outlier from Esperstedt before it – it is more likely a recent migrant from the steppe. The most likely origin of this individual is therefore people from the steppe, i.e. either the Srubna culture or a related group. Its relatively close cluster in PCA to certain recent Slavic populations can be interpreted in light of the multiple back and forth migrations in the region: of steppe populations to the west (Srubna, Cimmerians, Scythians, Sarmatians,…), and of Slavic-speaking populations:
And then rapidly expanding as a Proto-Slavic-speaking community from the steppe to the west.
Well-defined outliers are, therefore, essential to understand a recent history of admixture. On the other hand, the very concept of “outlier” can be a dangerous tool – when the lack of enough samples makes their classification as as such unjustified -, leading to the wrong interpretations.
A preprint article by two of the most prolific researchers in Human Ancestry is out, and they request feedback: Ancient genomics: a new view into human prehistory and evolution, by Skoglund and Mathieson (2017). Right now, it is downloadable on Dropbox.
The first decade of ancient genomics has revolutionized the study of human prehistory and evolution. We review new insights based on ancient genomic data, including greatly increased resolution of the timing and structure of the out-of-Africa event, the diversification of present-day non-African populations, and the earliest expansions of those populations into Eurasia and America. Prehistoric genomes now document patterns of population continuity and change on every inhabited continent–in particular the effect of agricultural expansions in Africa, Europe and Oceania–and record a history of natural selection that shapes present-day phenotypic diversity. Despite these advances, much remains unknown, in particular about the genomic histories of Asia–the most populous continent, and Africa–the continent that contains the most genetic diversity. Ancient genomes from these and other regions, integrated with a growing understanding of the genomic basis of human phenotypic diversity, will be in focus during the next decade of research in the field.
The paper may be highly recommended as an introduction for anyone interested in the field of Human Ancestry in general.
The next substantial change is closely related to ancestry that by around 5000 BP extended over a region of more than 2000 miles of the Eurasian steppe, including in individuals associated with the Yamnaya Cultural Complex in far-eastern Europe (1; 38) and with the Afanasievo culture in the central Asian Altai mountains (1). This “steppe” ancestry is itself a mixture between ancestry that is related to Mesolithic hunter-gatherers of eastern Europe and ancestry that is related to both present-day populations (38) and Mesolithic hunter-gatherers (46) from the Caucasus mountains, and also to the populations of Neolithic (11), and Copper Age (56) Iran. Steppe ancestry appeared in southeastern Europe by 6000 BP (72), northeastern Europe around 5000 BP (47) and central Europe at the time of the Corded Ware Complex around 4600 BP (1; 38). These dates are reasonably tight constraints, because in each case there is no evidence of steppe ancestry in individuals immediately preceding these dates (47; 72). Gene flow on the steppe was extensive and bidirectional, as shown by the eastward flow of Anatolian Neolithic ancestry– reaching well into central Eurasia by the time of the Andronovo culture ~3500 BP (1)–and the westward flow of East Asian ancestry–found in individuals associated with the Iron Age Scythian culture close to the Black Sea ~2500 BP (143).
Copper and Bronze Age population movements (14; 78 Martiniano, 2017 #8761; 85; 112), as well as later movements in the Iron Age and Historical period (70; 119) further distributed steppe ancestry around Europe. Present-day western European populations can be modeled as mixtures of these three ancestry components (Mesolithic hunter-gatherer, Anatolian Neolithic and Steppe) (38; 57). In eastern Europe, further shifts in ancestry are the result of additional or distinct gene flow from Anatolia throughout the Neolithic and Bronze Age in the Aegean (42; 51; 55; 72; 87), and gene flow from Siberian-related populations in Finland and the Baltic region (38). East-west gene flow also brought new ancestry–related to populations from 265 Copper Age Iran–to the Levant during the Copper and Bronze ages (39; 56).
The geographic structure of these population transformations gave rise to population structure of present-day Europe. For example Anatolian Neolithic ancestry is highest in southern European populations like Sardinians, and lowest in northern European populations (38). Steppe ancestry is at high frequency in north-central Europeans and low in the south. Isolation-by-distance may have contributed to these patterns to some extent, but the contribution must have been small. In much of Europe, extreme population discontinuity was the norm.
Featured image: from the article, “Major Holocene population movements and expansions that have been demonstrated using ancient DNA.”
After my first version, findings in Olalde et al. (2017) and Mathieson et al. (2017) supported some of my predictions. Now after my third, their new data also supports another prediction. Because the model is based on solid linguistic and archaeological models. Here is an excerpt from the Indo-European demic diffusion model, 3rd ed. (pp. 55-56):
At the end of the Trypillian culture, herding/hunting trends intensified, and the agricultural system collapsed, with people moving to the steppe zone, as confirmed by the presence of numerous graves to the south (Rassamakin 1999). At the same time, the Trypillian world absorbed a foreign tradition related to materials of settlement sites of the Dnieper steppes – such as the late Sredni Stog culture –, like cord impressions and burial rites similar to the later Corded Ware culture, marking also the transformation of decors and changes in their interpretation (Palaguta 2007).
The similarity in burial rituals between Yamna and Corded Ware made Gimbutas define a common “Kurgan people”, whose relationship has also been long supported by Kristiansen (Kristiansen 1989; Kristiansen et al. 2017). An equivalence of both burial rites has been, however, rejected (Häusler 1963, 1978, 1983), and it is generally agreed that the Yamna culture did not expand to the north of the Tisza River.
The importance of horse exploitation in Deriivka, in the forest-steppe zone of the north Pontic region along the Dnieper region, during the Middle Eneolithic period (probably ca. 3700-3530 BC), suggests that horses played a significant role in the life of this Sredni Stog community (Anthony and Brown 2003). In its late period (ca. 4000-3500 BC), this culture had adopted corded ware pottery, and stone battle-axes.
However, this [sic] western steppe peoples were mainly hunters (Rassamakin 1999), and the ‘herding skill’ essential for wild horse domestication seems absent (Kuzmina 2003). All this has been confirmed with zooarchaeological evidence and new molecular and stable isotope results, suggesting an absence of horse domestication in territories of the late Sredni Stog culture in the north Pontic steppe (Mileto et al. 2017), before the advent of migrants from the Indo-European-speaking Repin culture.
The new sample described in Mathieson et al. (2017), dated ca. 4200 BC (but within a wide range, 5000-3500 BC) is from a site classified as of late Sredni Stog (although potentially from Post-Mariupol / Kvitjana), a culture of hunters who probably did not breed domesticated horses (even after the period of conquest and dominance of Suvorovo-Novodanilovka chiefs, from Indo-Hittite-speaking early Khvalynsk, who had domesticated horses), and – more importantly – is of R1a-M417 lineage, shows high so-called “Yamna component” in ADMIXTURE, and clusters among Corded Ware samples in PCA approximately a thousand years before this culture’s expansion. Information from the supplementary material:
An Eneolithic cemetery of the Sredny Stog II culture was excavated by D. Telegin in 1955-1957 near the village of Alexandria, Kupyansk district, Kharkov region on the left bank of the river Oskol. A total of 33 individuals were recovered. Based on craniometric analysis (I.Potekhina 1999) it was suggested that the Eneolithic inhabitants of Alexandria were not homogeneous and resulted from admixture of local Neolithic hunter-gatherers and early farmers, possibly Trypillian groups. We report genetic data from one individual: I6561
Another individual from Eneolithic Ukraine (of R1b1 xM269 lineage) clusters quite closely with Neolithic samples from the Baltic, which points to the strong connection between both – southern and northern – regions of east-central Europe before the period of great Chalcolithic expansions, and the potential origin of the spread of R1b (xM269) lineages with the Corded Ware culture.
It will be fun to see the mess that certain researchers have made (and will still make in the near future) of their findings coupled with the concept of “Yamna component”, when trying to describe the “proxy ancestral populations” of European Copper Age and Bronze Age cultures… Difficult times ahead for many, after the collapse of the simplistic Yamna -> Corded Ware -> Bell Beaker genetic model laid out since Haak et al. (2015) and Allentoft et al. (2015).
[EDIT 27 September 2017] Not directly related, but here is today’s interesting discussion on Twitter surrounding the ancestral populations of the “Yamnaya component”, for illustration of the discussions to come when this ancestry is divided into different, more precise, older (Neolithic) steppe components, and these in turn shown to contribute to different European and Asian Chalcolithic and Bronze Age cultures:
Rough attempt to understand genetic history of Europe and W. Eurasia. It's tough to get your head around. Comments welcome. pic.twitter.com/lutXFKmluk
Given the variance found in the three samples from Eneolithic Ukraine (comparable to the variance found in east Bell Beaker samples), we may now be getting closer to the precise territory and culture where the Corded Ware culture might have formed, which cannot be much further from the Dnieper-Dniester region before the Yamna expansion to the west ca. 3300 BC, judging from the elevated steppe component.
It seems, because of the proximity of both cultures and the similar dates of their migrations, that the westward expansion of the Yamna culture may have indeed provided an important push (among some strong ‘pull’ forces) for peoples of the expansion of the Corded Ware culture.
So Genetics reinforces the solidest models of Archaeology and Linguistics? Professional academics being mostly right in their careful research, and amateur geneticists playing with software being wrong? Who would have thought… More and more papers help thus shut up naysayers who state (again and again) that new algorithms are here to revolutionise these academic fields.
The expansion of peoples is known to be associated with the spread of a certain admixture component + the expansion and reduction in variability of a haplogroup (i.e. few male lineages are usually more successful during the expansion): Neolithic farmers from the Middle East expanding with haplogroup G2a; Natufian component (Levant hunter-gatherers or later, Neolithic farmers) and haplogroup E southward into Africa; CHG component expansion with haplogroup J; WHG expansion into east Europe with haplogroup R1b; etc.
There were (at least) two main expansion processes involving Proto-Indo-European: one causing the branching off of the language ancestral to Anatolian, and another during the spread of Late Indo-European dialects. Based on this, and on known archaeological models, I have predicted since the first version of the demic diffusion model:
Based on haplogroups found until then in Yamna (R1b-M269), Corded Ware (R1a-M417, especially Z645), and Bell Beaker (R1b-L151):
that mainly R1b-L23 (especially L51) lineages and more steppe admixture would be found in east Bell Beaker – confirmed some two months after my publication by Olalde et al. (2017);
and that mainly R1a-M417 (especially Z645) subclades will be found in Corded Ware samples.
Based on the finding of “Yamna component” in the Corded Ware culture: that this admixture must have come from somewhere else. I pointed out to eastern Europe, including the forest and forest-steppe zone especially in the natural continuum of the Dniester-Dnieper region. Especially after Mathieson et al. (2017), in my second and third versions of the model, I have more specifically suggested a southern origin in the region, nearer to where the CHG ancestry must have come from (the Caucasus and cultures formed in contact with it), according to mainstream archaeological data, i.e. cultures of the North Pontic steppe / steppe-forest. But of course, until more samples are available, more CHG ancestry in other cultures of the Forest Zone cannot be discarded.
For the vast majority of academics, more samples (regionally proportioned) are needed only from early Corded Ware, as we have from Bell Beaker: if they are (as expected) mostly R1a-M417, then everything is clear, and it will finally mean the end for the tiring, now almost ‘traditional’ association R1a – Proto-Indo-European. Some more samples from the potential homeland of the third Corded Ware horizon, most likely Ukraine (Podolia and Volynia regions), nearer to the time of the Corded Ware expansion, would also be great, to locate the actual ancestral population of Corded Ware migrants – recognisable by the main presence of haplogroup R1a-Z645 (formed ca. 3500 BC), and elevated “Yamna component” before the arrival of the Yamna culture…
If, however, early Corded Ware samples of R1b-L23 subclades are found in certain quantity, especially old samples from east-central Europe (excluding Yamna migrants along the Prut), the tricky question of Late Indo-European cultural diffusion will remain: Did Corded Ware peoples adopt a Late Indo-European language from clans of R1b-L23 lineages? That is what Kristiansen and Anthony have been betting for, a cultural diffusion, caused by:
A long-lasting contact, according to Kristiansen (1989,…,2017). He defends that Sredni Stog adopted the language – but obviously not the same culture – from the east, but that it is a genetic and cultural mix from Globular Amphora, Trypillia, and steppe cultures. This has been Kristiansen’s model for almost 30 years, and it follows Marija Gimbutas’ outdated theory of the “Kurgan people”.
A rapid change according to Anthony (2007). He associates the adoption of Pre-Germanic with the domination of Yamna chiefs over Usatovo people, and the adoption of Balto-Slavic by the people from (Corded Ware) Middle Dnieper group because of the technical superiority of neighbouring Yamna herders.
Linguistics, with the growing support of a North-West Indo-European group, points clearly to a European expansion of a community speaking the ancestral language of Italo-Celtic, Germanic, and probably Balto-Slavic. Archaeology, too, showed migration from Yamna only to south-eastern Europe (correcting Gimbutas’ Kurgan model) and later with east Bell Beaker mainly into central, western, and northern Europe.
Even Kristiansen admits that only after the arrival of Bell Beaker in Scandinavia was a linguistic community (i.e. Germanic) formed – although he places the center of gravity in Úněticean influence, and (yet again) a cultural diffusion event into the Danish Dagger period.
Because of more and more data contrasting with old theories, some have elected to develop weak, indemonstrable links, to keep supporting e.g. Gimbutas’ concept of “Kurgan people” in Archaeology, and a sudden, early expansion of all PIE dialects at once in Linguistics. It seems that, after so much fuss about the (misleading) ‘Yamna component’ concept – and so many far-fetched assumptions by amateur geneticists -, the Corded Ware connection will once again hinge on weak, indemonstrable cultural diffusion theories, be it ‘Kurgan peoples’ (including now, of course, Eneolithic cultures of Ukraine) or any culture from eastern Europe that will reveal some close samples to Corded Ware migrants, in terms of PCA, ADMIXTURE, or haplogroup.
So once we find mainly R1a-Z645 in more Corded Ware samples (and this haplogroup and more “Yamna component” in non-Yamna cultures of Eneolithic Ukraine, and potentially Poland or Belarus) we all may finally expect a peaceful acceptance of reality, at least in Genetics? Nope. No siree. Nein. Not then, not ever.
Why? Because some people want their paternal lineage to have lived in their historical region, and spoken their historical language, since time immemorial. It won’t matter if Archaeology, Linguistics, Genetics, etc. don’t support their claims: if they need to use some aspects of admixture, or haplogroups (or a combination of them) from carefully selected samples instead of looking at the whole picture; if they have to support that Indo-Europeans came from a culture different than Yamna, in- or outside of the steppe or forest-steppe, be it the Balkans, Anatolia, Armenia, or the Moon; if their proto-language should then come directly from Indo-Hittite, or from a Germano-Slavonic, or Indo-Slavonic, or Indo-Germanic group, or whatever invented dialectal branch necessary to fit their model, or if they have to support the ‘constellation analogy’ of Clackson, or thousands of years of development for each branch; etc. They will support whatever is necessary.
And this adaptation, obviously, has no end. It’s stupid, I know. But that’s how we are, how we think. We have seen that these sad trends continue no matter what, for decades, and not only regarding Indo-European. Some common examples include:
Indo-Aryan-speaking Indians defending an autochthonous origin of R1a and Indo-European; as well as the ‘opposite’ autochtonous continuity theory of Dravidian-speaking Indians (based on ASI ancestry, haplogroup R2, mtDNA haplogroup M, or whatever is at hand).
Western Europeans defending an autochthonous origin of the R1b haplogroup, with a Palaeolithic or Mesolithic origin, including the language, viz. the recent Indo-European from the Atlantic façade theories (in the Celtic from the West series, by Koch and Cunliffe); the now fading Palaeolithic Continuity Theory; and many other forgotten Eurocentric proposals; as well as the more recent informal hints of a central European/Balkan homeland based on the Villabruna cluster and south-eastern Mesolithic finds, which is at risk of being related to a Balkan origin of Proto-Indo-European…
There is also the ‘opposite’ theory of the autochthonous origin of the Basques, including Proto-Iberians and potentially other peoples like Paleo-Sardinians, based on the previously popular Vasconic-Uralic hypothesis (and an ancient Europe divided into R1b and N1c1 haplogroups), which is still widely believed in certain regions.
Nordic speakers supporting the autochthonous nature of Germanic and haplogroup I1 to Scandinavia.
Armenian speakers delighted to see a proposal of Indo-European homeland in the Armenian highlands, be it supported by glottalic consonants, CHG ancestrty, R1b (xM269) or J lineages…
Greek speakers now willing to support continuity of haplogroup J as a ‘native’ Greek lineage, of people speaking Proto-Greek (and in earlier times PIE), because of two Minoan, and one Mycenaean samples found in Lazaridis et al. (2017).
Even Turks linking Yamna with the expansion of Turkic languages. That one is fun to read, almost like a parody for the rest – substituting “Indo-European” for “Turkic”.
For years, a lot of people – me included (at least since 2005) – believed, because of modern maps of R1a distribution, that R1a and Corded Ware are the vector of Indo-European languages. For those of us who don’t have any personal or national tie with this haplogroup, this notion has been easy to change with new data. For others, it obviously isn’t, and it won’t be.
For all these people, a sample, result, or conclusion from any paper, just dubiously in favour, means everything, but a thousand against mean nothing, or can be reinterpreted to support their fantasies.
The Kossinnian “autochthonous continuity” crap permeates this relatively new subfield of Human Evolutionary Genetics, as it permeated Indo-European studies (first Linguistics, then Archaeology) in its infancy. It seems to be a generalised human trend, no doubt related to some absurd inferiority complex, mixed with historical romanticism, a certain degree of chauvinism, and (falling in the eternal Godwin’s Law of our field) some outdated, childish notion of ‘supremacy’ linked with the expansion of the own language and people.
Such simplistic and popular models are also lucrative, judging by the boom in demand for DNA analysis, which companies embellish with modern fortune tellers (or fortune tellers themselves sell for a price), promising to ascertain your ‘ancestry proportions’ using automated algorithms, so that you don’t have to get lost in complex genetic data and prehistoric accounts, which can’t help you define your “ethnicity”…
Some just don’t want to realize that the spread of prehistoric languages (like Late Indo-European dialects) was a complex, non-uniform, stepped process, devoid of modern romantic concepts, which in genetic terms necessarily included later founder effects and cultural diffusions, so that no one can trace their haplogroup, lineage, family, region, or country to any single culture, language, or ethnic group. The same, by the way, can be said of peoples and countries in historic times.
As I said before, we shall expect supporters of the Kurgan model (and thus the expansion of R1a-Z645 with Yamna) to wait for just one sample of R1a-M417 in Yamna and/or Bell Beaker (which will eventually be found), and just one sample of R1b-M269 in Corded Ware (which will also eventually be found), to blow the horn of victory in this naïve competition against time, general knowledge, and (essentially) themselves.
A sad consequence of how we are is that, because of the obvious influence of these stupid modern ethnolinguistic agendas, because we are not all rowing in the same direction, genetic results and conclusions are still perceived as far-fetched and labile, and thus most archaeologists and linguists prefer not to include genetic results in their investigation. And those who dare to do so, are badly counselled by those who go with the tide, so that their papers become almost instantly outdated.