Corded Ware and Bell Beaker related groups defined by patrilocality and female exogamy

tumulus-culture-eba-danube

Two new interesting papers concerning Corded Ware and Bell Beaker peoples appeared last week, supporting yet again what is already well-known since 2015 about West Uralic and North-West Indo-European speakers and their expansion.

Below are relevant excerpts (emphasis mine) and comments.

NOTE. I will add analyses of ancestry, renewed Y-DNA maps, etc. if and when I find the time.

I. Corded Ware and Battle Axe cultures

Open access The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon, by Malmström, Günther, et al. Philos. Trans. R. Soc. (2019).

I.1. Origins of Corded Ware peoples

The discovery of the Alexandria outlier represented a clear support for a long-lasting genomic difference between the two distinct cultural groups, Yamnaya and Corded Ware, already visible in an opposition Khvalynsk vs. late Sredni Stog ca. 4000 BC, i.e. well before the formation of both Late Eneolithic/Early Bronze Age groups.

However, the realization that it may not have been an Eneolithic individual, but rather a (Middle?) Bronze Age one, suggests that Sredni Stog was possibly not directly related to Corded Ware, and a potential direct connection with Yamnaya might have to be reevaluated, e.g. through the Carpathian Basin, as Anthony (2017) proposed.

pca-yamnaya-corded-ware-oblaczkowo
Principal component analysis of modern Europeans (grey) and projected ancient Europeans.

This new paper shows two early Corded Ware individuals from Obłaczkowo, Poland (ca. 2900-2600 BC) – hence close to the supposed original Proto-Corded Ware community – with an apparently (almost) full “Steppe-like” ancestry, clustering (almost) with Yamnaya individuals:

Similar to the BAC individuals, the newly sequenced individuals from the present-day Karlova in Estonia and Obłaczkowo in Poland appear to have strong genetic affinities to other individuals from BAC and CWC contexts across the Baltic Sea region. Some individuals from CWC contexts, including the two from Obłaczkowo, cluster closely with the potential source population of steppe-related ancestry, the Yamnaya herders. Notably, these individuals appear to be those with the earliest radiocarbon dates among all genetically investigated individuals from CWC contexts. Overall, for CWC-associated individuals, there is a clear trend of decreasing affinity to Yamnaya herders with time.

NOTE. Interestingly, this sample is almost certainly attributed to the skeleton E8-A, which had been supposedly already investigated by the Copenhagen group as the RISE1 sample:

We note that RISE1 is also described as the individual from Obłaczkowo feature E8-A. However, their genetic results differ from ours. They present this individual as a molecularly determined male that belongs to Y-chromosomal haplogroup (hg) R1b and to mtDNA hg K1b1a1 while our results show this individual to be female, carrying a mtDNA hg U3a’c profile

Since the typical Steppe_MLBA ancestry of Corded Ware groups does not show good fits for (Pre-)Yamnaya-derived ancestry, it is almost certain that these individuals will show no (or almost no) direct Yamnaya-related contribution, but rather a contribution of East European sub-Neolithic groups, more or less close to the steppe-forest region.

NOTE. They might show contributions from Pre-Yamnaya-influenced Sredni Stog, though, but if they show a contribution of Yamnaya, then they are probably outliers, related to Yamnaya vanguard groups (see image below). And for them to show it, then both sources, Yamnaya and Corded Ware, should be clearly distinguishable from each other and their relative contribution quantifiable in formal stats, something difficult (if not impossible) to ascertain today.

Their position in the published PCA – a plot apparently affected by projection bias – suggests a cluster in common with early Baltic samples, which are known to show contributions from East European sub-Neolithic populations (see qpAdm values for Baltic CWC samples).

NOTE. Results for previous samples labelled as Poland CWC are unreliable due to their low coverage.

The most interesting aspect about the ancestry shown by these early samples is their further support for an origin of the culture different than Sredni Stog, and for a rejection of the Alexandria outlier as ancestral to them, hence for a Volhynian-Podolian homeland of Proto-Corded Ware peoples, with an ancestry probably more closely related to the late Maykop Steppe- and Trypillian/GAC groups admixed with sub-Neolithic populations of the Eastern European Late Eneolithic.

NOTE. That is, unless there is a reason for the apparent increase in so-called “Steppe-ancestry” during the northward and westward migration of CWC peoples that represents another thing entirely…

trypillian-yamnaya-influence-baltic
Trypillian routes of influence and Yamnaya culture influences in Central and Central-East Europe during the Late Eneolithic / Early Bronze Age. Images by Klochko (2009).

I.2. CWC expansion under R1a bottlenecks

The two males in our dataset (ber1 and poz81) belonged to Y-chromosome R1a haplogroups, as do the majority of males (16/24) from the previously published CWC contexts, while a smaller fraction belonged to R1b [3/24] or I2a [3/24] lineages. The R1a haplogroup has not been found among Neolithic farmer populations nor in hunter–gatherer groups in central and western Europe, but it has been reported from eastern European hunter–gatherers and Eneolithic groups. Individuals from the Pontic–Caspian steppe, associated with the Yamnaya Culture, carry mostly R1b and not R1a haplotypes.

Sample poz81 is of basal hg. R1a-CTS4385*, an R1a-M417 subclade, supporting once again that most Corded Ware individuals from western and central European groups expanded under R1a-M417 (xZ645) lineages. The Battle Axe sample from Bergsgraven (ca. 2620-2470 BC) shows a basal hg. R1a-Y2395*, a R1a-Z283 subclade leading to the typically Fennoscandian R1a-Z284.

Both findings further support that typical lineages of West CWC groups, including R1a-M417 (xZ645) subclades, were fully replaced by incoming East Bell Beakers, and that the limited expansion of R1a-Z284 and I1 (the latter found in one newly reported Late Neolithic sample from Sweden) was the outcome of later regional bottlenecks within Scandinavia, after the creation of a maritime dominion by the Bell Beaker elites during the Dagger Period.

I.3. CWC and lactase persistence

(…) one of these individuals (kar1) carried at least one allele (-13910 C->T) associated with lactose tolerance, while the other two individuals (ber1 and poz81) carried at least one ancestral variant each, consistent with previous observations of low levels of lactose tolerance variants in the Neolithic and a slight increase among individuals from CWC contexts.

The fact that two early CWC individuals carry ancestral variants could be said to support the improbability of the individual from Alexandria representing a community ancestral to the Corded Ware community. On the other hand, the late CWC individual from Estonia carries one allele, but it still seems that only Bell Beakers and Steppe-related groups show the necessary two alleles during the Early Bronze Age, which is in line with a late Repin/early Yamnaya-related origin of the successful selection of the trait, consistent with the expansion of their specialized semi-nomadic cattle-breeding economy through the steppe biome during the Late Eneolithic.

rs4988235-lactase-persistence-history
Maps part of the public data used for the post by Iain Mathieson on Lactase Persistence. “By 2500 BP, the allele is present over a band stretching from Ireland to Central Asia at around 50 degrees latitude. This probably reflects the spread of Steppe ancestry populations in which the allele originated. However, the allele is still rare (say less than 1% frequency) over this entire range. It does not become common anywhere until some time in the past 2500 years – when it reaches its present-day high frequency in Britain and Central Europe”.

I.4. West Uralic spread from the East

The BAC groups fit as a sister group to the CWC-associated group from Estonia but not as a sister group to the CWC groups from Poland or Lithuania (|Z| > 3), indicating some differences in ancestry between these CWC groups and BAC. Supervised admixture modelling suggests that BAC may be the CWC-related group with the lowest YAM-related ancestry and with more ancestry from European Neolithic groups.

While the results of the paper are compatible with a migration from either the Eastern or the Western Baltic into Scandinavia, phylogeography and archaeology support that Battle Axe peoples emerged as a Baltic Corded Ware group close to the Vistula that expanded first to the north-east, and then to the west from Finland, continuing mostly unscathed during the whole Bronze Age mostly in eastern Fennoscandia with the development of Balto-Finnic- and Samic-speaking communities.

corded-ware-culture-ancestry-over-time
Correlation between f4(Chimp, LBK, YAM, X), where X is a CWC or BAC individual, and the date (BCE) of each individual. This statistic measures shared drift between CWC and Linear Pottery Culture (LBK) as opposed to YAM and should increase with the higher proportion of Neolithic farmer ancestry in CWC and BAC.

Radiocarbon dating showed that the three individuals from the Öllsjö megalithic tomb derived from later burials, where oll007 (2860–2500 cal BCE) overlaps with the time interval of the BAC, and oll009 and oll010 (1930–1650 cal BCE) fall within the Scandinavian Late Neolithic and Early Bronze Age

For more on how the Pitted Ware culture may have influenced Uralic-speaking Battle Axe peoples earlier than Indo-European-speaking Bell Beakers in Scandinavia, read more about Early Bronze Age Scandinavia and about the emergence of the Pre-Proto-Germanic community.

II. Bell Beakers through the Bronze Age

New paper (behind paywall) Kinship-based social inequality in Bronze Age Europe, by Mittnik et al. Science (2019).

II.1. Yamnaya vanguard settlers

In my last post, I showed how the ancestry of Corded Ware from Esperstedt is consistent with influence by incoming Yamnaya vanguard settlers or early Bell Beakers, stemming ultimately from the Carpathian Basin, something that could be inferred from the position of the Esperstedt outlier in the PCA, and by the knowledge of Yamnaya archaeological influences up to Saxony-Anhalt.

Yamnaya settlers are strongly suspected to have migrated in small so-called vanguard groups to the west and north of the Carpathians in the first half of the 3rd millennium BC, well before the eventual adoption of the Proto-Beaker package and their expansion ca. 2500 BC as East Bell Beakers.

Tauber Valley infiltration

As I mentioned in the books, one of the known – among the many more unknown – sites displaying Yamnaya-related traits and suggesting the expansion of Yamnaya settlers into Central Europe is Lauda-Königshofen, in the Tauber Valley.

From Diet and Mobility in the Corded Ware of Central Europe, by Sjögren, Price, & Kristiansen PLoS One (2017):

A series of CW cemeteries have been excavated in the Tauber valley. There are three large cemeteries known and some 30 smaller sites. The larger ones are Tauberbischofsheim-Dittingheim with 62 individuals, Tauberbischofsheim-Impfingen with 40 individuals, and Lauda-Königshofen with 91 individuals. The cemeteries are dispersed rather regularly along the Tauber valley, on both sides of the river, suggesting a quite densely settled landscape.

The Lauda-Königshofen graves consisted mostly of single inhumations in contracted position, usually oriented E-W or NE-SW. A total of 91 individuals were buried in 69 graves. At least 9 double graves and three graves with 3–4 individuals were present. In contrast to the common CW pattern, sexes were not distinguished by body position, only by grave goods. This trait is common in the Tauber valley and suggests a local burial tradition in this area. Stone axes were restricted to males, pottery to females, while other artifacts were common to both sexes. About a third of the graves were surrounded by ring ditches, suggesting palisade enclosures and possibly over-plowed barrows.

In particular, Frînculeasa, Preda, & Heyd (2015) used Lauda-Königshofen as representative of the mobility of horse-riding Yamnaya nomadic herders migrating into southern Germany, referring to the findings in Trautmann (2012) about the nomadic herders from the Tauber Valley, and their already known differences with other Corded Ware groups.

The likely influence of Yamnaya in the region has been reported at least since the 2000s, repeatedly mentioned by Jozef Bátora (2002, 2003, 2006), who compiled Yamnaya influences in a map that has been copied ever since, with little improvement over time. Heyd believes that there are potentially many Yamnaya remains along the Middle and Lower Danube and tributaries not yet found, though.

NOTE. Looking for this specific site, I realized that Bátora (and possibly many after him who, like me, copied his map) located Lauda-Königshofen in a more south-western position within Baden-Württemberg than its actual location. I have now corrected it in the maps of Chalcolithic migrations.

yamnaya-corded-ware-europe
Yamnaya influences in Central Europe suggestive of vanguard settlements, contemporary with Corded Ware groups. See full map.

Althäuser Hockergrab…Bell Beakers

Unfortunately, though, it is very difficult to attribute the reported R1b-L51 sample from the Tauber valley to a population preceding the arrival of East Bell Beakers in the region, so there is no uncontroversial smoking gun of Yamnaya vanguard settlers – yet. Reasons to doubt a Pre-Beaker origin are as follows:

1. This family of the Tauber valley shows a late radiocarbon date (ca. 2500 BC), i.e. from a time where East Bell Beakers are known to have been already expanding in all directions from the Middle and Upper Danube and its tributaries.

tauber-valley-althauser-hockergrab
Crouched burial from Althausen (Althäuser Hockergrab), dated ca. 2500 BC.

2. Archaeological information is scarce. Remains of these four individuals were discovered in 1939 and officially reported together with other findings in 1950, without any meaningful data that could distinguish between Bell Beakers and Corded Ware individuals.

This site is located in the Tauber valley, ca. 100 km to the northwest of the Lech valley. The site was discovered during the construction of a sports field in 1939 and was subsequently excavated by G. Müller and O. Paret. Four individuals in crouched position were found in the burial pit of a flat grave. The burial did not contain any grave goods, but due to the type of grave and positioning of the bodies (with heads pointing towards southwest) the site was attributed to the Corded Ware complex.

The classification of this burial as of CWC and not BBC seems to have been based entirely on the numerous CWC findings in the Tauber valley, rather than on its particular burial orientation following a regional custom (foreign to the described standard of both cultures), and on its grave type that was also found among Bell Beaker groups. Like many human remains recovered in dubious circumstances in the 20th century, these samples should have probably been labelled (at least in the genetic paper) more properly as Tauber_LN or Tauber_EBA.

yamnaya-bias-tauber-lech-valley
Changes in ancestry over time. (A) Median ages of individuals plotted against z scores of f4 (Mbuti, Test; Yamnaya_Samara, Anatolia_Neolithic) show increase of Anatolian farmer-related ancestry (indicated by more positive z-scores) and decrease of variation in ancestry over time. Grey shading indicates significant z scores, red line shonw near correlation (r = -0.35971; P = 0.003) and dotted lines the 95% confidence interval. (B) ancestry proportions on autosomes calculated with qpAdm. (C) Sex-bias z scores between autosomes and X chromosomes show significant male bias for steppe-related ancestry in the Tauber samples. Image modified from the paper: Surrounded with a blue circle in (A) are females with more Steppe-related ancestry, and in (C) surrounded by squares are the distinct sex biases found in the earliest BBC from the Tauber valley vs. later groups from the Lech valley.

3. In terms of ancestry, there seem to be no gross differences between the Lech Valley BBC individuals and previously reported South German Beakers, originally Yamnaya-like settlers admixing through exogamy with locals, including Corded Ware peoples, as the sex bias of the Lech Valley Beakers proves (see PCA plot below). In other words, northern and eastern Beakers admixed with regional (Epi-)Corded Ware females during their respective expansions, similar to how southern and western Beakers admixed with regional EEF-related females.

The two available Tauber Valley samples (“Tauber_CWC”) show the same pattern: a quite recent Steppe-related male bias and Anatolia_Neolithic-related female bias. Nevertheless, the male sample clusters ‘to the south’ in the PCA relative to all sampled Corded Ware individuals (see PCA plot below), and shows less Yamnaya-like ancestry than what is reported (or can be inferred) for Yamnaya from Hungary or early Bell Beakers of elevated Steppe-related ancestry.

The ancestry and position of the Althäuser male in the PCA is thus fully compatible with recently incoming East Bell Beakers admixing with local peoples (including Corded Ware) through exogamy, but not so much with a sample that would be expected from Yamanaya vanguard + Corded Ware-related ancestry (more like the Esperstedt outlier or the early France Beaker). Compared to the more ‘northern’ (fully Corded Ware-like) position of his female counterpart, there is little to support that both are part of the same native Tauber valley community after generations of ancestry levelling…

yamnaya-ancestry-tauber-cwc-bbc-lech-eba-mba
Table S9. Three-way qpAdm admixture model for European MN/Chalcolithic group+Yamnaya_Samara. P-values greater than 0.05 (model is not rejected) marked in green.

4. The haplogroup inference is also unrevealing: whereas the paper reports that it is R1b-P310* (xU106, xP312), there is no data to support a xP312 call, so it may well be even within the P312 branch, like most sampled Bell Beaker males. Similarly, the paper also reports that HUGO_180Sk1 (ca. 2340 BC) shows a positive SNP for the U106 trunk, which would make it the earliest known U106 sample and originally from Central Europe, but there is no clear support for this SNP call, either. At least not in their downloadable BAM files, as far as I can tell. Even if both were true, they would merely confirm the path of expansion of Yamnaya / East Bell Beakers through the Danube, already visible in confirmed genomic data:

r1b-l51-archaic-yamnaya-bell-beakers
Distribution of ‘archaic’ R1b-L51 subclades in ancient samples, overlaid over a map of Yamnaya and Bell Beaker migrations. In blue, Yamnaya Pre-L51 from Lopatino (not shown) and R1b-L52* from BBC Augsburg. In violet, R1b-L51 (xP312,xU106) from BBC Prague and Poland. In maroon, hg. R1b-L151* from BBC Hungary, BA Bohemia, and (not shown) a potential sample from the Tauber Valley and one from BBC at Mondelange, which is certainly xU106, maybe xP312. Interestingly, the earliest sample of hg. R1b-U106 (a lineage more proper of northern Europe) has been found in a Bell Beaker from Radovesice (ca. 2350 BC), between two of these ‘archaic’ R1b-L51 samples; and a sample possibly of hg. R1b-ZZ11+ (ancestral to DF27 and U152) was found in a Bell Beaker from Quedlinburg, Germany (ca. 2290 BC), to the north-west of Bohemia. The oldest R1b-U152 are logically from Central Europe, too.

II.2. Proto-Celts and the Tumulus culture

The most interesting data from Mittnik et al. (2019) – overshadowed by the (at first sight) striking “CWC” label of the Althäuser male – is the finding that the most likely (Pre-)Proto-Celtic community of Southern Germany shows, as expected, major genetic continuity over time with Yamnaya/East Bell Beaker-derived patrilineal families, which suggests an almost full replacement of other Y-chromosome haplogroups in Southern German Bronze Age communities, too.

Sampled families form part of an evolving Bell Beaker-derived European BA cluster in common with other Indo-European-speaking cultures from Western, Southern, and Northern Europe, also including early Balto-Slavs, clearly distinct from the Corded Ware-related clusters surviving in the Eastern Baltic and the forest zone.

This Central European Bronze Age continuity is particularly visible in many generations of different patrilocal families practising female exogamy, showing patrilineal inheritance mainly under R1b-P312 (mostly U152+) lineages proper of Central European bottlenecks, all of them apparently following a similar sociopolitical system spanning roughly a thousand years, since the arrival of East Bell Beakers in the region (ca. 2500 BC) until – at least – the end of the Middle Bronze Age (ca. 1300 BC):

Here, we show a different kind of social inequality in prehistory, i.e., complex households that consisted of i) a higher-status core family, passing on wealth and status to descendants, ii) unrelated, wealthy and high-status non-local women and iii) local, low-status individuals. Based on comparisons of grave goods, several of the high-status non-local females could have come from areas inhabited by the Unetice culture, i.e., from a distance of at least 350 km. As the EBA evidence from most of Southern Germany is very similar to the Lech valley, we suggest that social structures comparable to our microregion existed in a much broader area. The EBA households in the Lech valley, however, seem similar to the later historically known oikos, the household sphere of classic Greece, as well as the Roman familia, both comprising the kin-related family and their slaves.

pca-lech-valley-bell-beaker-eba
Genetic structure of Late Neolithic and Bronze Age individuals from southern Germany. (A) Ancient individuals (covered at 20,000 or more SNPs) projected onto principal components defined by 1129 present day west Eurasians (shown in fig. S6); individuals in this study shown with outlines corresponding to their 87Sr/86Sr isotope value (black: consistent with local values, orange: uncertain/intermediate, red: inconsistent with local values). Selected published ancient European individuals are shown without outlines. Image modified from the paper. Surrounded by triangles in cyan, Corded Ware-like females; with a blue triangle, Yamnaya/Early BBC-like sample from the Tauber valley.

NOTE. For those unfamiliar with the usual clusters formed by the different populations in the PCA, you can check similar graphics: PCA with Bell Beaker communities, PCA with Yamnaya settlers from the Carpathians, a similar one from Wang et al. (2019) showing the Yamnaya-Hungary cline, or the chronological PCAs prepared by me for the books.

The gradual increase in local EEF-like ancestry among South Germany EBA and MBA communities over the previous BBC period offers a reasonable explanation as to how Italic and Celtic communities remained in loose contact (enough to share certain innovations) despite their physical separation by the Alps during the Early Bronze Age, and probably why sampled Bell Beakers from France were found to be the closest source of Celts arriving in Iberia during the Urnfield period.

Furthermore, continued contacts with Únětice-related peoples through exogamy also show how Celtic-speaking communities closer to the Danube might have influenced (and might have been influenced by) Germanic-speaking communities of the Nordic Late Neolithic and Bronze Age, helping explain their potentially long-lasting linguistic exchange.

Like other previous Neolithic or Chalcolithic groups that Yamnaya and Bell Beakers encountered in Europe, ancestry related to the Corded Ware culture became part of Bell Beaker groups during their expansion and later during the ancestry levelling in the European Early Bronze Age, which helps us distinguish the evolution of Indo-European-speaking communities in Europe, and suggests likely contacts between different cultural groups separated hundreds of km. from each other.

All in all, there is nothing to support that (epi-)Corded Ware groups might have survived in any way in Central or Western Europe: whether through their culture, their Y-chromosome haplogroups, or their ancestry, they followed the fate of other rapidly expanding groups before them, viz. Funnelbeaker, Baden, or Globular Amphorae cultural groups. This is very much unlike the West Uralic-speaking territory in the Eastern Baltic and the Russian forests, where Corded Ware-related cultures thrived during the Bronze Age.

lech-valley-yamnaya-ancestry-over-time
f4-statistics showing differences in ancestry in populations grouped by period. An increase in affinity to ancestry related to Anatolia Neolithic over time. Males and females grouped together shown as upward and downward pointing triangles, respectively.

Conclusion

It was about time that geneticists caught up with the relevance of Y-DNA bottlenecks when assessing migrations and cultural developments.

From Malmström et al. (2019):

The paternal lineages found in the BAC/CWC individuals remain enigmatic. The majority of individuals from CWC contexts that have been genetically investigated this far for the Y-chromosome belong to Y-haplogroup R1a, while the majority of sequenced individuals of the presumed source population of Yamnaya steppe herders belong to R1b. R1a has been found in Mesolithic and Neolithic Ukraine. This opens the possibility that the Yamnaya and CWC complexes may have been structured in terms of paternal lineages—possibly due to patrilineal inheritance systems in the societies — and that genetic studies have not yet targeted the direct sources of the expansions into central and northern Europe.

From Gibbons (2019), a commentary to Mittnik et al. (2019):

Some of the early farmers studied were part of the Neolithic Bell Beaker culture, named for the shape of their pots. Later generations of Bronze Age men who retained Bell Beaker DNA were high-ranking, buried with bronze and copper daggers, axes, and chisels. Those men carried a Y chromosome variant that is still common today in Europe. In contrast, low-ranking men without grave goods had different Y chromosomes, showing a different ancestry on their fathers’ side, and suggesting that men with Bell Beaker ancestry were richer and had more sons, whose genes persist to the present.

There was no sign of these women’s daughters in the burials, suggesting they, too, were sent away for marriage, in a pattern that persisted for 700 years. The only local women were girls from high-status families who died before ages 15 to 17, and poor, unrelated women without grave goods, probably servants, Mittnik says. Strontium levels from three men, in contrast, showed that although they had left the valley as teens, they returned as adults.

Also, from Scientific American:

(…) it has long been assumed that prior to the Athenian and Roman empires,—which arose nearly 2,500 and more than 2,000 years ago, respectively—human social structure was relatively straightforward: you had those who were in power and those who were not. A study published Thursday in Science suggests it was not that simple. As far back as 4,000 years ago, at the beginning of the Bronze Age and long before Julius Caesar presided over the Forum, human families of varying status levels had quite intimate relationships. Elites lived together with those of lower social classes and women who migrated in from outside communities. It appears early human societies operated in a complex, class-based system that propagated through generations.

It seems wrong (to me, at least) that the author and – as he believes – archaeologists and historians had “assumed” a different social system for the European Bronze Age, which means they hadn’t read about how Indo-European societies were structured. For example, long ago Benveniste (1969) already drew some coherent picture of these prehistoric peoples based on their reconstructed language alone: regarding their patrilocal and patrilineal family system; regarding their customs of female exogamy and marriage system; and regarding the status of foreigners and slaves as movable property in their society.

A long-lasting and pervasive social system of Bronze Age elites under Yamnaya lineages strikingly similar to this Southern German region can be easily assumed for the British Isles and Iberia, and it is likely to be also found in the Low Countries, Northern Germany, Denmark, Italy, France, Bohemia and Moravia, etc., but also (with some nuances) in Southern Scandinavia and Central-East Europe during the Bronze Age.

Therefore, only the modern genetic pool of some border North-West Indo-European-speaking communities of Europe need further information to describe a precise chain of events before their eventual expansion in more recent times:

  1. the relative geographical isolation causing the visible regional founder effects in Scandinavia, proper of the maritime dominion of the Nordic Late Neolithic (related thus to the Island Biogeography Theory); and
  2. the situation of the (Pre-)Proto-Balto-Slavic community close to the Western Baltic which, I imagine, will be shown to be related to a resurge of local lineages, possibly due to a shift of power structures similar to the case described for Babia Góra.

NOTE. Rumour has it that R1b-L23 lineages have already been found among Mycenaeans, while they haven’t been found among sampled early West European Corded Ware groups, so the westward expansion of Indo-European-speaking Yamnaya-derived peoples mainly with R1b-L23 lineages through the Danube Basin merely lacks official confirmation.

Related

Bell Beakers and Mycenaeans from Yamnaya; Corded Ware from the forest steppe

eba-yamnaya-ancestry-hungary

I have recently written about the spread of Pre-Yamnaya or Yamnaya ancestry and Corded Ware-related ancestry throughout Eurasia, using exclusively analyses published by professional geneticists, and filling in the gaps and contradictory data with the most reasonable interpretations. I did so consciously, to avoid any suspicion that I was interspersing my own data or cherry picking results.

Now I’m finished recapitulating the known public data, and the only way forward is the assessment of these populations using the available datasets and free tools.

Understanding the complexities of qpAdm is fairly difficult without a proper genetic and statistical background, which I won’t pretend to have, so its tweaking to get strictly correct results would require an unending game of trial and error. I have sadly little time for this, even taking my tendency to procrastination into account… so I have used a simple model akin to those published before – in particular, the outgroup selection by Ning, Wang et al. (2019), who seem to be part of the only group interested in distinguishing Yamnaya-related from Corded Ware-related ancestry, probably the most relevant question discussed today in population genomics regarding the Proto-Indo-European and Proto-Uralic homelands.

eneolithic-steppe-best-fits
Supplementary Table 13. P values of rank=2 and admixture proportions in modelling Steppe ancestry populations as a three-way admixture of Eneolithic steppe Anatolian_Neolithic and WHG using 14 outgroups.
Left populations: Test, Eneolithic_steppe, Anatolian_Neolithic, WHG.
Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.

I have used for all analyses below a merged dataset including the curated one of the Reich Lab, the latest on Central and South Asia by Narasimhan, Patterson et al. (2019), on Iberia by Olalde et al. (2019), and on the East Baltic by Saag et al. (2019), as well as datasets including samples from Wang et al. (2019) and Lamnidis et al. (2018). I used (and intend to use) the same merged dataset in all cases, despite its huge size, to avoid adding one more uncontrolled variable to the analyses, so that all results obtained can be compared.

I try to prepare in advance a bunch of relevant files with left pops and right pops for each model:

  1. It seems a priori more reasonable to use geographically and chronologically closer proxy populations (say, Trypillia or GAC for Steppe-related peoples) than hypothetic combinations of ancestral ones (viz. Anatolian farmer, WHG, and EHG).
  2. This also means using subgroups closer to the most likely source population, such as (Don-Volga interfluve) Yamnaya_Kalmykia rather than (Middle Volga) Yamnaya_Samara for the western expansion of late Repin/early Yamnaya, or the early Germany_Corded_Ware.SG or Czech_Corded Ware for the group closest to the Proto-Corded Ware population (see below), likely neighbouring the Upper Vistula region.
  3. I usually test two source populations for different targets, which seems like a much more efficient way of using computer resources, whenever I know what I want to test, since I need my PC back for its normal use; whenever I don’t know exactly what to test, I use three-way admixture models and look for subsets to try and improve the results.

I have probably left out some more complex models by individualizing the most relevant groups, but for the time being this would have to do. Also, no other formal stats have been used in any case, which is an evident shortcoming, ruling out an interpretation drawn directly and only from the results below.

Full qpAdm results for each batch of samples are presented in a Google Spreadsheet, with each tab (bottom of the page) showing a different combination of sources, usually in order of formally ‘best’ (first to the left) to ‘worst’ (last to the right) fits, although the order is difficult to select in highly heterogeneous target groups, as will be readily visible.

maykop-trypillia-intrusion-steppes
Disintegration, migration, and imports of the Azov–Black Sea region. First migration event (solid arrows): Gordineşti–Maikop expansion (groups: I – Bursuchensk; II – Zhyvotylivka; III – Vovchans’k; IV – Crimean; V – Lower Don; VI – pre-Kuban). Second migration event (hollow arrows): Repin expansion. After Rassamakin (1999), Demchenko (2016).

Corded Ware origins

The latest publications on the Yampil barrow complex have not improved much our understanding of the complexity of Corded Ware origins from an archaeological point of view, involving multiple cultural (hence likely population) influences. This bit is from Ivanova et al., Baltic-Pontic Studies (2015) 20:1, and most hypotheses of the paper remain unanswered (except maybe for the relevance of the Złota group):

In the light of the above outline therefore one should argue that the ‘architecture of barrows’ associated in the ‘Yampil landscape’ of the Middle Dniester Area with the Eneolithic (specifically, mainly with the TC), precedes the development of a similar phenomenon that can be observed from 2900/2800 BC in the Upper Dniester Area and drainage basin of the Upper Vistula, associated with the CWC [Goslar et al. 2015; Włodarczak 2006; 2007; 2008; Jarosz, Włodarczak 2007]. The most consuming research question therefore is whether ritual customs making use of Eneolithic (Tripolye) ‘barrow architecture’ could have penetrated northwards along the Dniester route, where GAC communities functioned. One could also ask what role the rituals played among the autochthons [Kośko 2000; Włodarczak 2008; 2014: 335; Ivanova, Toshchev 2015b].

This issue has already been discussed with a resulting tentative systemic taxonomy in the studies of Włodarczak, arguing for the Złota culture (ZC) in the Vistula region as an illustration of one of the (Małopolska) reception centres of civilization inspirations from the oldest Pontic ‘barrow culture’ circle associated with the Eneolithic and Early Bronze Age [Włodarczak 2008]. Notably, it is in the ZC that one can notice a set of cultural traits (catacomb grave construction, burial details, forms and decoration of vessels) analogous to those shared by the north-western Black Sea Coast groups of the forest-steppe Eneolithic (chiefly Zhyvotilovka-Volchansk) and the Late Tripolye circle (chiefly Usatovo-Gordinești-Horodiștea-Kasperovtsy).

gac-trypillia-usatovo-corded-ware
Globular Amphorae culture „exodus” to the Danube Delta: a – Globular Amphorae culture; b – GAC (1), Gorodsk (2), Vykhvatintsy (3) and Usatovo (4) groups of Trypillia culture; c – Coţofeni culture; d – northern border of the late phase of Baden culture;red arrows – direction of Globular Amphora culture expansion; blue arrow – direction of „reflux” of Globular Amphora culture (apud Włodarczak, 2008, with changes).

Taking into account that I6561 might be wrongly dated, we cannot include the Corded Ware-like sample of the end-5th millennium BC in the analysis of Corded Ware origins. That uncertainty in the chronology of the appearance of “Steppe ancestry” in Proto-Corded Ware peoples complicates the selection of any potential source population from the CHG cline.

Nevertheless, the lack of hg. R1a-M417 and sizeable Pre-Yamnaya-related ancestry in the sampled Pontic forest-steppe Eneolithic populations (represented exclusively by two samples from Dereivka ca. 3600-3400 BC) would leave open the interesting possibility that a similar ancestry got to the forest-steppe region between modern Poland and Ukraine during the known complex population movements of the Late Eneolithic.

It is known that Corded Ware-derived groups and Steppe Maykop show bad fits for Pre-Yamnaya/Yamnaya ancestry, and also that Steppe Maykop is a potential source of “Steppe-related ancestry” within the Eneolithic CHG mating network of the Pontic-Caspian steppes and forest-steppes. Testing Corded Ware for recent Trypillia and Maykop influences, proper of Late Trypillia and Late Maykop groups in the North Pontic area (such as Zhyvotylivka–Vovchans’k and Gordineşti) side by side with potential Pre-Yamnaya and Yamnaya sources makes thus sense:

Now, the main obvious difference between Khvalynsk-Yamnaya and Corded Ware is the long-lasting, pervasive Y-chromosome bottlenecks under R1b lineages in the former, compared to the haplogroup variability and late bottleneck under R1a-M417 in the latter, which speaks in favour – on top of everything else – of a different community of sub-Neolithic hunter-gatherers including hg. R1a-M417 hijacking the expansion of Steppe_Maykop-related ancestry around the Volhynian-Podolian Upland.

Akin to how Yamnaya patrilineal descendants hijacked regional EEF (±CWC) ancestry components mainly through exogamy, dragging them into the different expanding Bell Beaker groups (see below), but kept their Indo-European languages, these hunter-gatherers that admixed with peoples of “Steppe ancestry” were the most likely vector of expansion of Uralic languages in Eastern Europe.

corded-ware-from-trypillia-maykop
PCA of ancient Eurasian samples. Marked likely Proto-Corded Ware samples and potential origin of its PCA cluster based on qpAdm results. See full PCA and more related files.

Baltic Corded Ware

One of the most interesting aspects of the results above is the surprising heterogeneity of the different regional groups, which is also reflected in the Y-DNA variability of early Corded Ware samples.

Seeing how Baltic CWC groups, especially the early Latvia_LN sample, show particularly bad fits with the models above, it seems necessary to test how this population might have come to be. My first impression in 2017 was that they could represent early Corded Ware groups admixed with Yamnaya settlers through their interactions along the Dnieper-Dniester corridor.

However, I recently predicted that the most likely admixture leading to their ancestry and PCA cluster would involve a Corded Ware-like group and a group related to sub-Neolithic cultures of eastern Europe, whose best proxy to date are EHG-like Khvalynsk samples (i.e. excluding the outlier with Pre-Yamnaya ancestry, I0434):

corded-ware-pca-sub-neolithic-europe
Detail of the PCA of the Corded Ware expansion. See full PCA and more related files.

Late Corded Ware + Yamnaya vanguard

Relevant are also the mixtures of Corded Ware from Esperstedt, and particularly those of the sample I0104, which I have repeated many times in this blog I suspected to be influenced by vanguard Yamnaya settlers:

The infeasible models of CWC + Yamnaya_Kalmykia ± Hungary_Baden (see below for Bell Beakers) and the potential cluster formed with other samples from the Baltic suggest that it could represent a more complex set of mixtures with sub-Neolithic populations. On the other hand, its location in Germany, late date (ca. 2500 BC or later), and position in the PCA, together with the good fits obtained for Germany_Beaker as a source, suggest that the increase in Steppe-related ancestry + EEF makes it impossible for the model (as I set it) to directly include Yamnaya_Kalmykia, despite this excess Steppe-related ancestry actually coming from Yamnaya vanguard groups.

I think it is very likely that the future publication of EEF-admixed Yamnaya_Hungary samples (or maybe even Yamnaya vanguard samples) will improve the fits of this model.

These results confirm at least the need to distrust the common interpretation of mixtures including late Corded Ware samples from Esperstedt (giving rise to the “up to 75% Yamnaya ancestry of CWC” in the 2015 papers) as representative of the Corded Ware culture as a whole, and to keep always in mind that an admixture of European BA groups including Corded Ware Esperstedt as a source also includes East BBC-like ancestry, unless proven otherwise.

yamnaya-vanguard-corded-ware-chalcolithic-early
Yamnaya vanguard groups in Corded Ware territory before the expansion of Bell Beakers (ca. 2500 BC). See full map.

Bell Beaker expansion

A hotly (re)debated topic in the past 6 months or so, and for all the wrong reasons, is the origin of the Bell Beaker folk. Archaeology, linguistics, and different Y-chromosome bottlenecks clearly indicate that Bell Beakers were at the origin of the North-West Indo-European expansion in Europe, while the survival of Corded Ware-related groups in north-eastern Europe is clearly related to the expansion of Uralic languages.

NOTE. For the interesting case of Proto-Indo-Iranians expanding with Corded Ware-like ancestry, see more on the formation of Sintashta-Potapovka-Filatovka from East Uralic-speaking Abashevo and Pre-Proto-Indo-Iranian-speaking Poltavka herders. See also more on R1a in Indo-Iranians and on the social complexity of Sintashta.

Nevertheless, every single discarded theory out there seems to keep coming back to life from time to time, and a new wave of interest in “Bell Beaker from the Single Grave culture” somehow got revived in the process, too, because this obsession – unlike the “Bell Beakers from Iberia Chalcolithic” – is apparently acceptable in certain circles, for some reason.

We know that Iberian Beakers, British Beakers, or Sicilian EBA – representing the most likely closest source population of speakers of Proto-Galaico-Lusitanian, Pre-Celtic Indo-European, and Proto-Elymian, respectively – have already been successfully tested for a direct origin among Western European Beakers in Olalde et al. (2018), Olalde et al. (2019), and Fernandes et al. (2019).

This success in ascertaining a closer Beaker source is probably due to the physical isolation of the specific groups (related to Germany_Beaker, Netherlands_Beaker, and NE_Mediterranean_Beaker samples, respectively) after their migration into regions dominated by peoples without Steppe-related ancestry. Furthermore, Celtic-speaking populations expanding with Urnfield south of the Pyrenees also show a good fit with a source close to France_Beaker.

So I decided to test sampled Bell Beaker populations, to see if it could shed light to the most likely source population of individual Beaker groups and the direction of migration within Central Europe, i.e. roughly eastwards or westwards. As it was to be expected for closely related populations (see the relevant discussion here), an attempt to offer a simplistic analysis of direction based on formal stats does not make any sense, because most of the alternative hypotheses cannot be rejected:

Not only because of the similar values obtained, but because it is absurd to take p-values as a measure of anything, especially when most of these conflicting groups with slightly ‘better’ or ‘worse’ p-values represent multiple different mixtures of the type (Yamnaya + EEF) + (Corded Ware + EEF ± Yamnaya), impossible to distinguish without selecting proper, direct ancestral populations…

A further example of how explosive the Bell Beaker expansion was into different territories, and of their extensive local admixture, is shown by the unsuccessful attempt by Olalde et al. (2018) to obtain an origin of the EEF source for all Beaker groups (excluding Iberian Beakers):

bell-beaker-local-population-iberia
Investigating the genetic makeup of Beaker-complex-associated individuals. Testing different populations as a source for the Neolithic ancestry component in Beaker-complex-associated individuals. The table shows P values (* indicates values > 0.05) for the fit of the model: ‘Steppe_EBA + Neolithic/Copper Age’ source population.
burials-yamnaya-hungary
Map of attested Yamnaya pit-grave burials in the Hungarian plains; superimposed in shades of blue are common areas covered by floods before the extensive controls imposed in the 19th century; in orange, cumulative thickness of sand, unfavourable loamy sand layer. Marked are settlements/findings of Boleráz (ca. 3500 BC on), Baden (until ca. 2800 BC), Kostolac (precise dates unknown), and Yamna kurgans (from ca. 3100/3000 BC on).

Now, there is a simpler way to understand what kind of Steppe-related ancestry is proper of Bell Beakers. I tested two simple models for some Beaker groups: Yamnaya + Hungary Baden vs. Corded Ware + GAC Poland. After all, the Bell Beaker folk should prefer a source more closely related to either Yamnaya Hungary or Central European Corded Ware:

Interestingly, models including Yamnaya + Baden show good fits for the most important groups related to North-West Indo-Europeans, including Bell Beakers from Germany, the Netherlands, Italy, and Poland, representing the most likely closest source populations of speakers of Pre-Proto-Celtic, Pre-Proto-Germanic, Proto-Italo-Venetic, and Pre-Proto-Balto-Slavic, respectively.

The admixed Yamnaya samples from Hungary that will hopefully be published soon by the Jena Lab will most likely further improve these fits, especially in combination with intermediate Chalcolithic populations of the Middle and Upper Danube and its tributaries, to a point where there will be an absolute chronological and geographical genomic trail from the fully Yamnaya-like Yamnaya settlers from Hungary to all North-West Indo-European-speaking groups of the Early Bronze Age.

The only difference between groups will be the gradual admixture events of their source Beaker group with local populations on their expansion paths, including peoples of mainly EEF, CWC+EEF, or CWC+EEF+Yamnaya related ancestry. There is ample evidence beyond ancestry models to support this, in particular continued Y-DNA bottlenecks under typical Yamnaya paternal lineages, mainly represented by R1b-L51 subclades.

east-bell-beaker-group-expansion
Distribution of the Bell Beaker East Group, with its regional provinces, as of c. 2400 cal BC (after Heyd et al. 2004, modified). See full maps.

European Early Bronze Age

European EBA groups that might show conflicting results due to multiple admixture events with Corded Ware-related populations are the Únětice culture and the Nordic Late Neolithic.

The results for Únětice groups seem to be in line with what is expected of a Central European EBA population derived from Bell Beakers admixed with surrounding poulations of East Bell Beaker and/or late (Epi-)Corded Ware descent.

Potential models of mixture for Nordic Late Neolithic samples – despite the bad fits due to the lack of direct ancestral CWC and BBC groups from Denmark – seem to be impossible to justify as derived exclusively from Single Grave or (even less) from Battle Axe peoples, supporting immigration waves of Bell Beakers from the south and further admixture events with local groups through maritime domination.

PCA of ancient European samples. Marked are Bronze Age clusters. See full PCAs.

Balkans Bronze Age

The potential origin of the typical Corded Ware Steppe-related ancestry in the social upheaval and population movements of the Dnieper-Dniester forest-steppe corridor during the 4th millennium BC raises the question: how much do Balkan Bronze Age groups owe their ancestry to a population different than the spread of Pre-Yamnaya-like Suvorovo-Novodanilovka chieftains? Furthermore, which Bronze Age groups seem to be more likely derived exclusively from Pre-Yamnaya groups, and which are more likely to be derived from a mixture of Yamnaya and Pre-Yamnaya? Do the formal stats obtained correspond to the expected results for each group?

Since the expansion of hg. I2a-L699 (TMRCA ca. 5500 BC) need not be associated with Yamnaya, some of these values – together with the assessment of each individual archaeological culture – may question their origin in a Yamnaya-related expansion rather than in a Khvalynsk-related one.

NOTE. These are the last ones I was able to test yesterday, and I have not thought these models through, so feel free to propose other source and target groups. In particular, complex movements through the North Pontic area during the Late Eneolithic would suggest that there might have been different Steppe-ancestry-related vs. EEF-related interactions in the north-west and west Pontic area before and during the expansion of Yamnaya.

Mycenaeans

One of the key Indo-European populations that should be derived from Yamnaya to confirm the Steppe hypothesis, together with North-West Indo-Europeans, are Proto-Greeks, who will in turn improve our understanding of the preceding Palaeo-Balkan community. Unfortunately, we only have Mycenaean samples from the Aegean, with slight contributions of Steppe-related ancestry.

Still, analyses with potential source populations for this Steppe ancestry show that the Yamnaya outlier from Bulgaria is a good fit:

The comparison of all results makes it quite evident the why of the good fits from (Srubnaya-related) Bulgaria_MLBA I2163 or of Sintashta_MLBA relative to the only a priori reasonable Yamnaya and Catacomb sources: it is not about some hypothetical shared ancestor in Graeco-Aryan-speaking East Yamnaya– or even Catacomb-Poltavka-related groups, because all available Yamnaya-related peoples are almost indistinguishable from each other (at least with the sampling available today). These results reflect a sizeable contribution of similar EEF-related populations from around the Carpathians in both Steppe-related groups: Corded Ware and Yamnaya settlers from the Balkans.

mycenaeans-minyan-ware-greece-minoan
Cultural groups in and around the Balkans during the Early Bronze Age. See full maps.

qpAdm magic

In hobby ancestry magic, as in magic in general, it is not about getting dubious results out of thin air: misdirection is the key. A magician needs to draw the audience attention to ‘remarkable’ ancestry percentages coupled with ‘great’ (?) p-values that purportedly “prove” what the audience expects to see, distracting everyone from the true interesting aspects, like statistical design, the data used (and its shortcomings), other opposing models, a comparison of values, a proper interpretation…you name it.

I reckon – based on the examples above – that the following problems lie at the core of bad uses of qpAdm:

  1. In the formal aspect, the poor understanding of what p-values and other formal stats obtained actually mean, and – more importantly – what they don’t mean. The simplistic trend to accept results of a few analyses at face value is necessarily wrong, in so far as there is often no proper reasoning of what is being assessed and how, and there is never a previous opinion about what could be expected if the alternative hypotheses were true.
  2. In the interpretation aspect, the poor judgement of accompanying any results with simplistic, superficial, irrelevant, and often plainly wrong archaeological or linguistic data selected a posteriori; the inclusion of some racial or sociopolitical overtones in the mixture to set a propitious mood in the target audience; and a sort of ritualistic theatrics with the main theme of ‘winning’, that is best completed with ad hominems.

If you get rid of all this, the most reasonable interpretation of the output of a model proposed and tested should be similar to Nick Patterson’s words in his explanation of qpWave and qpAdm use:

Here we see that, at least in this analysis there are reasonable models with CordedWareNeolithic is a mix of either WHG or LBKNeolithic and YamnayaEBA. (…) The point of this note is not to give a serious phylogenetic analysis but the results here certainly support a major Steppe contribution to the Corded Ware population, which is entirely concordant with the archaeology [?].

Very far, as you can see, from the childish “Eureka! I proved the source!”-kind of thinking common among hobbyists.

The Mycenaean case is an illustrative example: if the Yamnaya outlier from Bulgaria were not available, and if one were not careful when designing and assessing those mixture models, the interpretation would range from erroneous (viz. a Graeco-Aryan substrate, as I initially thought) to impossible (say, inventing migration waves of Sintashta or Srubnaya peoples into Crete). The models presented above show that a contribution of Yamnaya to Mycenaeans couldn’t be rejected, and this alone should have been enough to accept Yamnaya as the most likely source population of “Steppe ancestry” in Proto-Greeks, pending intermediate samples from the Balkans. In other words, one could actually find that ‘the best’ p-values for source populations of Mycenaeans is a combination of modern Poles + Turks, despite the impracticality of such a model…

I haven’t been able to reproduce results which supposedly showed that Corded Ware is more likely to be derived from (Pre-)Yamnaya than other source population, or that Corded Ware is better suited as the ancestral population of Bell Beakers. The analyses above show values in line with what has been published in recent scientific papers, and what should be expected based on linguistics and archaeology. So I’ll go out on a limb here and say that it’s only through a careful selection of outgroups and samples tested, and of as few compared models as possible, that you could eventually get this kind of results and interpretation, if at all.

Whether that kind of special care for outgroups and samples is about (a) an acceptable fine-tuning of the analyses, (b) a simplistic selection dragged from the first papers published and applied indiscriminately to all models, or (c) cherry picking analyses until results fit the expected outcome, is a question that will become mostly irrelevant when future publications continue to support an origin of the expansion of ancient Indo-European languages in Khvalynsk- and Yamnaya-related migrations.

Feel free to suggest (reasonable) modifications to correct some of these models in the comments. Also, be sure to check out other values such as proportions, SD or SNPs of the different results that I might have not taken into account when assessing ‘good’ or ‘bad’ fits.

Related

Baltic Finns in the Bronze Age, of hg. R1a-Z283 and Corded Ware ancestry

estonian-bronze-age-dna

Open access The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East, by Saag et al. Current Biology (2019).

Interesting excerpts:

In this study, we present new genomic data from Estonian Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). The cultural background of stone-cist graves indicates strong connections both to the west and the east [20, 21]. The Iron Age (IA) tarands have been proposed to mirror “houses of the dead” found among Uralic peoples of the Volga-Kama region [22].

(…) The 33 individuals included 15 from EstBA, 6 from EstIA, 5 from Pre-Roman to Roman Iron Age Ingria (500 BC–450 AD) (IngIA), and 7 from Middle Age Estonia (1200–1600 AD) (EstMA) and yielded endogenous DNA ∼4%–88%, average genomic coverages ∼0.017–0.734×, and contamination estimates <4% (Table S1). We analyzed the data in the context of modern and other ancient individuals, including from Neolithic Estonia [13].

estonian-y-dna-bronze-iron-age
Archaeological Information, Genetic Sex, mtDNA and Y Chromosome Haplogroups, and Average Coverage of the Individuals of This Study. Modified from the paper to mark distinct Y-DNA haplogroups in the LBA and IA.

We identified chrY hgs for 30 male individuals (Tables 1 and S2; STAR Methods). All 16 successfully haplogrouped EstBA males belonged to hg R1a, showing no change from the CWC period, when this was also the only chrY lineage detected in the Eastern Baltic [11, 13, 30, 31]. Three EstIA and two IngIA individuals also belonged to hg R1a, but three EstIA males belonged to hg N3a, the earliest so far observed in the Eastern Baltic. Three EstMA individuals belonged to hg N3a, two to hg R1a, and one to hg J2b. ChrY lineages found in the Baltic Sea region before the CWC belong to hgs I, R1b, R1a5, and Q [10, 11, 12, 13, 17, 32]. Thus, it appears that these lineages were substantially replaced in the Eastern Baltic by hg R1a [10, 11, 12, 13], most likely through steppe migrations from the east [30, 31]. (…) Our results enable us to conclude that, although the expansion time for R1a1 and N3a3′5 in Eastern Europe is similar [25], hg N3a likely reached Estonia or at least became comparably frequent to modern Estonia [1] only during the BA-IA transition.

A clear shift toward West Eurasian hunter-gatherers is visible between European LN and BA (including Baltic CWC) and EstBA individuals, the latter clustering together with Latvian and Lithuanian BA individuals [11]. EstIA, IngIA, and EstMA individuals project between BA individuals and modern Estonians, partially overlapping with both.

(…) EstBA individuals are clearly distinguishable from Estonian CWC individuals as the former have more of the blue component most frequent in WHGs and less of the brown and yellow components maximized in Caucasus hunter-gatherers and modern Khanty, respectively. The individuals of EstBA, EstIA, IngIA, EstMA, and modern Estonia are quite similar to each other on average, indicating that the relatively high proportion of WHG ancestry in modern Eastern Baltic populations compared to other present-day Europeans [15] traces back to the BA.

estonian-pca-published
Detail of the PCA, modified from the paper to label populations. Estonian Bronze Age and Iron Age samples cluster close to Early Corded Ware from the Baltic.. Principal-component analysis results of modern West Eurasians with ancient individuals projected onto the first two components (PC1 and PC2). BA, Bronze Age; EF, early farmers; HG, hunter-gatherers; IA, Iron Age; IMA, Iron/Middle Ages; LN, Late Neolithic; LNBA, Late Neolithic/Bronze Age; MA, Middle Ages

When comparing Estonian CWC and EstBA using autosomal outgroup f3 and Patterson’s D statistics (Table S3), the latter is more similar to other Baltic BA populations, to Baltic IA and Middle Age (MA) populations, and also to populations similar to WHGs and Scandinavian hunter-gatherers (SHGs), but not to Estonian CCC (Figures 2A and S2A; Data S1). The increase in WHG or SHG ancestry could be connected to western influences seen in material culture [20, 21] and facilitated by a decline in local population after the CCC-CWC period [20]. A slight trend of bigger similarity of Estonian CWC to forest or steppe zone populations and of EstBA to European early farmer populations can also be seen.

(…) When comparing to modern populations, Estonian CWC is slightly more similar to Caucasus individuals but EstBA to Baltic populations and Finnic speakers (Figure 2B; Data S1). Outgroup f3 and D statistics do not reveal apparent differences when comparing EstBA to EstIA, EstIA to IngIA, and EstIA to EstMA (Data S1).

estonian-ba-ia-ancestry
qpAdm results. Error bars indicate one SE. Central MN, Central European Middle Neolithic; EstBA, Estonian Bronze Age; EstIA, Estonian Iron Age; IngIA, Ingrian Iron Age; EstMA, Estonian Middle Ages; WHG, western hunter-gatherers.

These results highlight how uniparental and autosomal data can lead to different demographic inferences—the genetic change between CWC and BA not seen in uniparental lineages is clear in autosomal data and the appearance of chrY hg N in the IA is not matched by a clear shift in autosomal profiles.

EstBA individuals have no Nganasan-related ancestry and EstIA, IngIA, and EstMA individuals on average have 2% or 4% (Figure 3; Data S1). The differentiation remains when using BA or IA Fennoscandian populations [26] instead of Nganasans (Data S1). Notably, the proportion of Nganasan-related ancestry varies between 0% and 12% among sampled EstIA, IngIA, and EstMA individuals (Data S1), which may suggest its relatively recent admixture into the target population. Moreover, two individuals from Kunda (0LS10 and V10) have the highest proportions of Nganasan ancestry among EstIA (6% and 8%), one of them has chrY hg N3a, and isotopic analysis suggests neither individual being born in Kunda [34].

About these two males from Tarand-graves, ‘foreign’ to Kunda:

0LS10: Male from tarand III (burial 9; TÜ 1325: L777), age 17–25 years [34]. He had a fragment of a sheep/goat bone and ceramics as grave goods. This burial has two radiocarbon dates: 2430 ± 35 BP (Poz-10801; 760–400 cal BC) and 2530 ± 41 BP (UBA-26114; 800–530 cal BC) [34]. According to the isotopic analysis, the person was not born in the vicinity of Kunda; his place of birth is still unknown (but south-western Finland and Sweden are excluded) [34]. Sampled tooth r P1.

V10: Male from tarand XI (burial 24; TÜ 1325: L1925), age 25–35 years [34], date 2484 ± 40 BP (UBA-26115; 790–430 cal BC) [34]. He had a few potsherds near the skull. Likewise, this person was not locally born [34]. Sampled tooth l P1.

estonia-bronze-iron-age-steppe-siberian
Autosomal Analyses’ Results for Gyvakarai1 as the closest available Corded Ware source for Balto-Finnic populations.

The paper shows thus:

  • Major continuity of ancestry from Corded Ware to modern Estonians, with only slight changes in different periods. In fact, one of the best fits for the Late Bronze Age ancestry is Gyvakarai1, one of the Corded Ware “outliers” described as “closer to Yamna”, which I already said may be closer to Sredni Stog/EHG populations instead. Another interesting take is that the change from Bronze Age to Iron Age corresponds to an increase in Baltic Corded Ware-related ancestry, rather than being driven by Siberian ancestry.
  • pca-mittnik-gyvakarai
    File modified by me from Mittnik et al. (2018) to include the approximate position of the most common ancestral components, and an identification of potential outliers. Zoomed-in version of the European Late Neolithic and Bronze Age samples. “Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). From Mittnik et al. (2018).
  • A Volosovo-related migration of hg. N1c with Netted Ware into the area seems to be discarded, based on the full replacement of paternal lines and continuity of R1a-Z283. It is only during the Tarand-grave period when a system of chiefdoms (spread from Ananyino/Akozino) brings haplogroup N1c to the Gulf of Finland. During the Iron Age, the proportion of paternal lineages is still clearly in favour of R1a (50% in the coast, 100% in Ostrobothnia), which indicates a gradual replacement led by elites, likely because of the incorporation of Akozino warrior-traders spreading all over the Baltic, bringing the described shared Mordvinic traits in Fennic.
  • finno-ugric-haplogroup-n
    Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).
  • The arrival of Akozino warrior-traders (bringing N1c and R1a lineages) was probably linked to this minimal “Nganasan-like” ancestry of some samples in the transition to the Iron Age. This arrival is supported by samples 0LS10 (the earliest hg. N1c) and V10 (of hg. R1a), both dated to ca. 800-400 BC, with V10 showing the highest “Nganasan-like” ancestry with 4.8%, both of them neighbouring samples showing 0%. This variable admixture among local and foreign paternal lineages might support the described social system of family alliances with intermarriages. In fact, a medieval sample, 0LS03_1 (hg. R1a) also shows a recent “Nganasan-like” ancestry, which probably points to the integration of different Arctic-related ancestry components among Modern Estonians, in this case related to Finnish expansions and thus integration of Levänluhta-related ancestry, as per the supplementary data.
  • NOTE. Such minimal proportions of “Nganasan-like” ancestry evidence the process of admixture of Volga Finns in Akozino territory through their close interactions with Permians of Ananyino, who in turn acquired this Palaeo-Arctic admixture most likely during the expansion of the linguistic community to hunter-gatherer territories, to the north of the Cis-Urals. This process of stepped infiltration and expansion without language change is not dissimilar to the one seen among Indo-Iranians and Balto-Slavs of hg. R1b, or Vasconic speakers of hg. I2a, although in the case of Baltic Finns of hg. R1a the process of infiltration and expansion of hg. N1c is much less dramatic, with no radical replacement anywhere before the huge bottlenecks observable in Finns.

  • The expansion of haplogroup N1c among Finnic populations, as we are going to see in samples from the Middle Ages such as Luistari, is the consequence of late founder effects after huge bottlenecks expected based on the analysis of modern populations. The expansion of N1c-VL29 is different in origin from that of N1c-Z1936 among Samic (later integrated into Finnish populations), most likely from the east and originally associated with Lovozero Ware.
haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders. Map from Ilumäe et al. (2016).

In spite of all this, the conclusion of the paper is (surprise!) that Siberian ancestry and hg. N heralded the arrival of Finnic to the Gulf of Finland in the Iron Age… However, this conclusion is supposedly* supported, not by their previous papers, but by a recent phylogenetic study by Honkola et al. (2013), which doesn’t actually argue for such a late ‘arrival’: it argues for the split of Balto-Finnic around 1500 BC.

NOTE. I say ‘supposedly’ because Kristiina Tambets, for example, has been following the link of Uralic with haplogroup N since the 2000s, so this is not some conclusion they just happened to misread from some random paper they Googled. In those initial assessments, she argued that the “ancient homeland” of the Tat C mutation suggested that Finno-Ugrians were in Fennoscandia before Indo-Europeans. Apparently, since haplogroup N appears later and from the east, it is now more important to follow this haplogroup than what is established in archaeology and linguistics.

Even in the referred paper, this split is considered an in situ development, since the phylogenetic study takes the information – among others – 1) from Parpola and Carpelan, who consider Netted Ware, a culture derived from Fatyanovo/Abashevo and Volosovo, as the culprit of the Finno-Ugric expansion; and 2) from Kallio (2006), who clearly states that Proto-Balto-Finnic (like Proto-Finno-Samic) was spoken around the Gulf of Finland during the Bronze Age. Both of them set the terminus ante quem of the language presence in the Baltic ca. 1900 BC.

Anyways, as a consequence of geneticists keeping these untenable pre-ancient DNA haplogroup-based arguments today, I expect to see this “Finnic” language expansion also described for the Western Baltic, Scandinavia or northern Europe, when this same proportion of hg. N1c and “Nganasan” ancestry is observed in Iron Age samples around the Baltic Sea. The nativist trends that this domination of “Finns” all over Northern Europe 2,500 years ago will create will be even more fun to read than the current ones…

EDIT (10 May 2019) How I see the reaction of many to ancient DNA, in keeping their old theories:

Related

Yekaterinovsky Cape, a link between the Samara culture and early Khvalynsk

ekaterinovsky-cape

We already had conflicting information about the elite individual from the Yekaterinovsky Cape and the materials of his grave, which seemed quite old:

For the burial of 45 in the laboratory of the University of Pennsylvania, a 14C date was obtained: PSUAMS-2880 (Sample ID 16068)> 30 kDa gelatin Russia. 12, Ekaterinovka Grave 45 14C age (BP) 6325 ± 25 δ 13C (‰) –23.6 δ15 N (‰) 14.5. The results of dating suggest chronological proximity with typologically close materials from Yasinovatsky and Nikolsky burial grounds (Telegini et al. 2001: 126). The date obtained also precedes the existing dates for the Khvalynsk culture (Morgunova 2009: 14–15), which, given the dominance of Mariupol traits of the burial rite and inventory, confirms its validity. However, the date obtained for human bones does not exclude the possibility of a “reservoir effect” when the age can increase three or more centuries (Shishlin et al. 2006: 135–140).

Now the same date is being confirmed by the latest study published on the site, by Korolev, Kochkina, and Stachenkov (2019) and it seems it is really going to be old. Abstract (in part the official one, in part newly translated for clarity):

For the first time, pottery of the Early Eneolithic burial ground Ekaterinovsky Cape is published. Ceramics were predominantly located on the sacrificial sites in the form of compact clusters of fragments. As a rule, such clusters were located above the burials, sometimes over the burials, some were sprinkled with ocher. The authors have identified more than 70 vessels, some of which have been partially reconstructed. Ceramic was made with inclusion of the crushed shell into molding mass. The rims of vessels had the thickened «collar»; the bottoms had a rounded shape. The ornament was located on the rims and the upper part of the potteries. Fully decorated vessels are rare. The vessels are ornamented with prints of comb and rope stamps, with small pits. A particularity of ceramics ornamentation is presented by the imprints of soft stamps (leather?) or traces of leather form for the making of vessels. The ornamentation, made up of «walking comb» and incised lines, was used rarely as well as the belts of pits made decoration under «collar» of a rim. Some features of the ceramics decoration under study relate it with ceramics of the Khvalynsk culture. The ceramics of Ekaterinovsky Cape burial ground is attributed by the authors to the Samara culture. The ceramic complex under study has proximity to the ceramics from Syezzhe burial ground and the ceramics of the second phase of Samara culture. The chronological position is determined by the authors as a later period than the ceramics from the Syezzhe burial ground, and earlier than the chronological position of ceramics of the Ivanovka stage of the Samara culture and the Khvalynsk culture.

ekaterinovsky-cape-pottery
Ceramics from Ekaterinovsky Cape burial ground. 1–2, 4–5, 7–11 – ceramics from aggregations; 3, 6 – ceramics from the cultural layer.

More specifically:

Based on ceramic fragments from a large vessel from a cluster of sq.m. 14, the date received was: SPb-2251–5673 ± 120 BP. The second date was obtained in fragments from the aggregation [see picture above] from the cluster of sq.m. 45–46: SPb-2252–6372 ± 100 BP. The difference in dating indicates that the process of determining the chronology of the burial ground is far from complete, although we note that the earlier date almost coincided with the date obtained from the human bone from individual 45 (Korolev, Kochkina, Stashenkov, 2018, p. 300).

Therefore, the ceramics of the burial ground Ekaterinovsky Cape possess an originality that determines the chronological position of the burial ground between the earliest materials of the burial type in Syezzhe and the Khvalynsk culture. Techno-typological features of dishes make it possible to attribute it to the Samara culture at the stage preceding the appearance of Ivanovska-Khvalynsk ceramics.

It seems that this site showed cultural influences from the upstream region near the Kama-Vyatka interfluve, too, according to Korolev, Kochkina, Stashenkov, and Khokhlov (2018):

In 2017, excavation of burial ground Ekaterinovsky Cape were continued, located in the area of the confl uence of the Bezenchuk River in the Volga River. During the new excavations, 14 burials were studied. The skeleton of the buried were in a position elongated on the back, less often – crooked on the back with knees bent at the knees. In one burial (No. 90), a special position of the skeleton was recorded. In the burial number 90 in the anatomical order, parts of the male skeleton. This gave grounds for the reconstruction of his original position in a semi-sitting position with the support of elbows on the bottom of the pit. Noteworthy inventory: on the pelvic bones on the left lay a bone spoon, near the right humerus, the pommel of a cruciform club was found. A conclusion is made about the high social status of the buried. The results of the analysis of the burial allow us to outline the closest circle of analogies in the materials of Khvalynsky I and Murzikhinsky burial grounds.

Important sites mentioned in both papers and in this text:

To sum up, it seems that the relative dates we have used until now have to be corrected: older Khvalynsk I Khvalynsk II individuals, supposedly dated ca. 5200-4000 BC (most likely after 4700 BC), and younger Yekaterinovsky individuals, supposedly of the fourth quarter of the 5th millennium (ca. 4250-4000 BC), are possibly to be considered, in fact, roughly reversed, if not chronologically, at least culturally speaking.

Interestingly, this gives a new perspective to the presence of a rare fish- or reptile-headed pommel-scepter, which would be natural in a variable period of expansion of the horse and horse-related symbolism, a cultural trait rooted in the Samara culture attested in Syezzhe before the unification of the symbol of power under the ubiquitous Khvalynsk-Suvorovo horse-headed scepters and related materials.

ekaterinovsky-cape-pommel-mace
Ekaterinovsky Cape Burial Ground. Inventory of the burial no 90: 1, 2 – stone pommel of the mace; 3, 4 – bone article.

The Khvalynsk chieftain

If the reported lineages from Yekaterinovsky Cape are within the R1b-P297 tree, but without further clades, as Yleaf comparisons may suggest, there is not much change to what we have, and R1b-M269 could actually represent a part of the local population, but also incomers from the south (e.g. the north Caspian steppe hunter-gatherers like Kairshak), the east (with hunter-gatherer pottery), or the west near the Don River (in contact with Mariupol-related cultures, as the authors inferred initially from material culture).

Just like R1a-M417 became incorporated into the Sredni Stog groups after the Novodanilovka-Suvorovo expansion, probably as incoming hunter-gatherer pottery groups from the north admixing with peoples of “Steppe ancestry”, R1b-M269 lineages might have expanded explosively only during the Repin expansion, and maybe (like R1b-L51 later) they formed just a tiny part of the clans that dominated the steppe during the Khvalynsk-Novodanilovka community.

On the other hand, the potential finding of various R1b-M269/L23 samples in Yekaterinovsky Cape (including an elite individual) would suggest now, as it was supported in the original report by Mathieson et al. (2015), that these ancient R1b lineages found in the Volga – Ural region are in fact most likely all R1b-M269 without enough coverage to obtain proper SNP calls, which would simplify the picture of Neolithic expansions (yet again). From the supplementary materials:

10122 / SVP35 (grave 12). Male (confirmed genetically), age 20-30, positioned on his back with raised knees, with 293 copper artifacts, mostly beads, amounting to 80% of the copper objects in the combined cemeteries of Khvalynsk I and II. Probably a high-status individual, his Y-chromosome haplotype, R1b1, also characterized the high-status individuals buried under kurgans in later Yamnaya graves in this region, so he could be regarded as a founder of an elite group of patrilineally related families. His MtDNA haplotype H2a1 is unique in the Samara series.

khvalynsk-cemetery
Khvalynsk cemetery and grave gifts. Grave 90 contained copper beads and rings, a harpoon, flint blades, and a bird-bone tube. Both graves (90 and 91) were partly covered by Sacrificial Deposit 4 with the bones from a horse, a sheep, and a cow. Center: grave goods from the Khvalynsk cemetery-copper rings and bracelets, polished stone mace heads, polished stone bracelet, Cardium shell ornaments, boars tusk chest ornaments, flint blades, and bifiacial projectile points. Bottom: shell-tempered pottery from the Khvalynsk cemetery. After Agapov, Vasiliev, and Pestrikova 1990; and Ryndina 1998, Figure 31. Modified from Anthony (2007).

This remarkable Khvalynsk chieftain, whose rich assemblage may correspond to the period of domination of the culture all over the Pontic-Caspian steppes, has been consistently reported as of hg. R1b-L754 in all publications, including Wang et al. (2018/2019) tentative SNP calls in the supplementary materials (obtained with Yleaf, as the infamous Narasimhan et al. 2018 samples), but has been variously reported by amateurs as within the R1b-M73, R1b-V88, or (lately) R1b-V1636 trees, which makes it unlikely that quality of the sample is allowing for a proper SNP call.

The fact that Mathieson et al. (2015) considered it a member of the R1b-M269 clans appearing later in Yamna seems on point right now, especially if samples from Yekaterinovka are all within this tree. The relevance of R1b-L23 in the expansion of Repin and Yamna is reminiscent of the influence of successful clans among Yamna offshoots, such as Bell Beakers, and among Bell Beaker offshoots during the Bronze Age all over Europe.

Taking these younger expansions as example, it seems quite likely based on cultural links that (at least part of) the main clans of Khvalynsk were of R1b-M269 lineage, stemming from a R1b-dominated Samara culture, in line with the known succeeding expansions and the expected strictly patriarcal and patrilineal society of Proto-Indo-Europeans, which would have exacerbated the usual reduction in Y-chromosome haplogroup variability that happens during population expansions, and the aversion towards foreign groups while the culture lasted.

pontic-steppe-neolithic
Cultures of the Pontic-Caspian steppes and forest-steppes and surrounding areas during the Neolithic.

The finding of R1b-L23 in Yekaterinovka, associated with the Samara culture, before or during the Khvalynsk expansion, and close to the Khvalynsk site, would make this Khvalynsk chieftain most likely a member of the M269 tree (paradoxically, the only R1b-L754 branch amateurs have not yet reported for it). Similarly, the sample of a “Samara hunter-gatherer” of Lebyazhinka, of hg. R1b-P297, could also be under this tree, just like most R1b-M269 from Yamna are downstream from R1b-L23, and most reported R1b-M269 or R1b-L23 from Bell Beakers are under R1b-L151.

On the other hand, we know of the shortcomings of attributing a haplogroup expansion to the best known rulers, such as the famous lineages previously wrongly attributed to Niall of the Nine Hostages or Genghis Khan. The known presence of R1b-V1636 up to modern Greeks would be in line with an ancient steppe expansion that we know will show up during the Neolithic, although it could also be a sign of a more recent migration from the Caucasus. The presence of a sister clade of R1b-L23, R1b-PF7562, among modern Balkan populations, may also be attributed to a pre-Yamna steppe expansion.

y-dna-khvalynsk
Y-DNA samples from Khvalynsk and neighbouring cultures. See full version here.

On SNP calls

I reckon that even informal reports on SNP calls, like any other analyses, should be offered in full: not only with a personal or automatic estimation of the result, but with a detailed explanation of the good, dubious, and bad calls, alternatives to that SNP estimation, and a motivated reasoning of why one branch should be preferred over others. Downloading a sample and giving an instruction using a free software tool is never enough, as it became crystal clear recently for the hilariously biased and flawed qpAdm reports on Dutch Bell Beakers as the ‘missing link’ between Corded Ware and Bell Beakers…

Another example I can recall is the report of a R1a-Z93 subclade in the R1a-M417 sample ca. 4000 BC from Alexandria, which seems rather unlikely, seeing how this subclade must have split and expanded explosively with R1a-Z645 to the east with eastern Corded Ware groups, i.e. 1,000 years later, just like Z282 lineages expanded mainly to the north-east. But then again, as with the Khvalynsk chieftain, I have only seen indirect reports of that supposed SNP (including Y26+!), so we should just stick with its officially reported R1a-M417 lineage. This upstream haplogroup was, in fact, repeated with Yleaf’s tentative estimates in Wang et al. (2019) supplementary materials…

The combination of inexperienced, biased, or simply careless design, analyses, and reports, including SNP calls and qpAdm analyses (whether in forums or publications), however well-intentioned (or not) they might be, are hindering a proper analysis of data, adding to the difficulties we already have due to the scarcity of samples, their limited coverage, and the lack of proper context.

Some people like to repeat ad nauseam that archaeology and/or linguistics are ‘not science’ whenever they don’t fit their beliefs and myths based on haplogroup and/or ancestry. But it’s becoming harder and harder to rely on certain genetic data, too, and on their infinite changing interpretations, much more than it is to rely on linguistic and archaeological research, including data, assessments, and discussions that are open for anyone to review…if one is truly interested in them.

Ancient Sardinia hints at Mesolithic spread of R1b-V88, and Western EEF-related expansion of Vasconic

nuragic-sardinia-neolithic

New preprint Population history from the Neolithic to present on the Mediterranean island of Sardinia: An ancient DNA perspective, by Marcus et al. bioRxiv (2019)

Interesting excerpts (emphasis mine, edited for clarity):

On the high frequency of R1b-V88

Our genome-wide data allowed us to assign Y haplogroups for 25 ancient Sardinian individuals. More than half of them consist of R1b-V88 (n=10) or I2-M223 (n=7).

Francalacci et al. (2013) identi fied three major Sardinia-specifi c founder clades based on present-day variation within the haplogroups I2-M26, G2-L91 and R1b-V88, and here we found each of those broader haplogroups in at least one ancient Sardinian individual. Two major present-day Sardinian haplogroups, R1b-M269 and E-M215, are absent.

Compared to other Neolithic and present-day European populations, the number of identi fied R1b-V88 carriers is relatively high.

(…)ancient Sardinian mtDNA haplotypes belong almost exclusively to macro-haplogroups HV (n = 16), JT (n = 17) and U (n = 9), a composition broadly similar to other European Neolithic populations.

r1b-v88-europe
Geographic and temporal distribution of R1b-V88 Y-haplotypes in ancient European samples. We plot the geographic position of all ancient samples inferred to carry R1b-V88 equivalent markers. Dates are given as years BCE (means of calibrated 2s radio-carbon dates). Multiple V88 individuals with similar geographic positions are vertically stacked. We additionally color-code the status of the R1b-V88 subclade R1b-V2197, which is found in most present-day African R1b-V88 carriers.

On the origin of a Vasconic-like Paleosardo with the Western EEF

(…) the Neolithic (and also later) ancient Sardinian individuals sit between early Neolithic Iberian and later Copper Age Iberian populations, roughly on an axis that differentiates WHG and EEF populations and embedded in a cluster that additionally includes Neolithic British individuals. This result is also evident in terms of absolute genetic differentiation, with low pairwise FST ~ 0.005 +- 0.002 between Neolithic Sardinian individuals and Neolithic western mainland European populations. Pairwise outgroup-f3 analysis shows a very similar pattern, with the highest values of f3 (i.e. most shared drift) being with Neolithic and Copper Age Iberia, gradually dropping off for temporally and geographically distant populations.

In explicit admixture models (using qpAdm, see Methods) the southern French Neolithic individuals (France-N) are the most consistent with being a single source for Neolithic Sardinia (p ~ 0:074 to reject the model of one population being the direct source of the other); followed by other populations associated with the western Mediterranean Neolithic Cardial Ware expansion.

sardinians-ancient-eef
Principal Components Analysis based on the Human Origins dataset. A: Projection of ancient individuals’ genotypes onto principal component axes de fined by modern Western Eurasians (gray labels).

Pervasive Western Hunter-Gatherer ancestry in Iberian/French/Sardinian population

Similar to western European Neolithic and central European Late Neolithic populations, ancient Sardinian individuals are shifted towards WHG individuals in the top two PCs relative to early Neolithic Anatolians Admixture analysis using qpAdm infers that ancient Sardinian individuals harbour HG ancestry (~ 17%) that is higher than early Neolithic mainland populations (including Iberia, ~ 8%), but lower than Copper Age Iberians (~ 25%) and about the same as Southern French Middle-Neolithic individuals (~ 21%).

sardinia-modern-ancient-nuragic-pca
Principal Components Analysis based on the Human Origins dataset. B: Zoom into the region most relevant for Sardinian individuals.

Continuity from Sardinia Neolithic through the Nuragic

We found several lines of evidence supporting genetic continuity from the Sardinian Neolithic into the Bronze Age and Nuragic times. Importantly, we observed low genetic differentiation between ancient Sardinian individuals from various time periods.

A qpAdm analysis, which is based on simultaneously testing f-statistics with a number of outgroups and adjusts for correlations, cannot reject a model of Neolithic Sardinian individuals being a direct predecessor of Nuragic Sardinian individuals (…) Our qpAdm analysis further shows that the WHG ancestry proportion, in a model of admixture with Neolithic Anatolia, remains stable at ~17% throughout three ancient time-periods.

sardinians-modern-ancient-pca-admixture
Present-day genetic structure in Sardinia reanalyzed with aDNA. A: Scatter plot of the rst two principal components trained on 1577 present-day individuals with grand-parental ancestry from Sardinia. Each individual is labeled with a location if at least 3 of the 4 grandparents were born in the same geographical location (\small” three letter abbreviations); otherwise with \x” or if grand-parental ancestry is missing with \?”. We calculated median PC values for each Sardinian province (large abbreviations). We also projected each ancient Sardinian individual on to the top two PCs (gray points). B/C: We plot f-statistics that test for admixture of modern Sardinian individuals (grouped into provinces) when using Nuragic Sardinian individuals as one source population. Uncertainty ranges depict one standard error (calculated from block bootstrap). Karitiana are used in the f-statistic calculation as a proxy for ANE/Steppe ancestry (Patterson et al., 2012).

Steppe influx in Modern Sardinians

While contemporary Sardinian individuals show the highest affinity towards EEF-associated populations among all of the modern populations, they also display membership with other clusters (Fig. 5). In contrast to ancient Sardinian individuals, present-day Sardinian individuals carry a modest “Steppe-like” ancestry component (but generally less than continental present-day European populations), and an appreciable broadly “eastern Mediterranean” ancestry component (also inferred at a high fraction in other present-day Mediterranean populations, such as Sicily and Greece).

Related

Iberia: East Bell Beakers spread Indo-European languages; Celts expanded later

iberia-migrations-celts

New paper (behind paywall), The genomic history of the Iberian Peninsula over the past 8000 years, by Olalde et al. Science (2019).

NOTE. Access to article from Reich Lab: main paper and supplementary materials.

Abstract:

We assembled genome-wide data from 271 ancient Iberians, of whom 176 are from the largely unsampled period after 2000 BCE, thereby providing a high-resolution time transect of the Iberian Peninsula. We document high genetic substructure between northwestern and southeastern hunter-gatherers before the spread of farming. We reveal sporadic contacts between Iberia and North Africa by ~2500 BCE and, by ~2000 BCE, the replacement of 40% of Iberia’s ancestry and nearly 100% of its Y-chromosomes by people with Steppe ancestry. We show that, in the Iron Age, Steppe ancestry had spread not only into Indo-European–speaking regions but also into non-Indo-European–speaking ones, and we reveal that present-day Basques are best described as a typical Iron Age population without the admixture events that later affected the rest of Iberia. Additionally, we document how, beginning at least in the Roman period, the ancestry of the peninsula was transformed by gene flow from North Africa and the eastern Mediterranean.

Interesting excerpts:

From the Bronze Age (~2200–900 BCE), we increase the available dataset (6, 7, 17) from 7 to 60 individuals and show how ancestry from the Pontic-Caspian steppe (Steppe ancestry) appeared throughout Iberia in this period (Fig. 1, C and D), albeit with less impact in the south (table S13). The earliest evidence is in 14 individuals dated to ~2500–2000 BCE who coexisted with local people without Steppe ancestry (Fig. 2B). These groups lived in close proximity and admixed to form the Bronze Age population after 2000 BCE with ~40% ancestry from incoming groups (Fig. 2B and fig. S6).

Y-chromosome turnover was even more pronounced (Fig. 2B), as the lineages common in Copper Age Iberia (I2, G2, and H) were almost completely replaced by one lineage, R1b-M269. These patterns point to a higher contribution of incoming males than females, also supported by a lower proportion of nonlocal ancestry on the X-chromosome (table S14 and fig. S7), a paradigm that can be exemplified by a Bronze Age tomb from Castillejo del Bonete containing a male with Steppe ancestry and a female with ancestry similar to Copper Age Iberians.

iberian-adna

For the Iron Age, we document a consistent trend of increased ancestry related to Northern and Central European populations with respect to the preceding Bronze Age (Figs. 1, C and D, and 2B). The increase was 10 to 19% (95% confidence intervals given here and in the percentages that follow) in 15 individuals along the Mediterranean coast where non-Indo-European Iberian languages were spoken; 11 to 31% in two individuals at the Tartessian site of La Angorrilla in the southwest with uncertain language attribution; and 28 to 43% in three individuals at La Hoya in the north where Indo-European Celtiberian languages were likely spoken (fig. S6 and tables S11 and S12).

This trend documents gene flow into Iberia during the Late Bronze Age or Early Iron Age, possibly associated with the introduction of the Urnfield tradition (18). Unlike in Central or Northern Europe, where Steppe ancestry likely marked the introduction of Indo-European languages (12), our results indicate that, in Iberia, increases in Steppe ancestry were not always accompanied by switches to Indo-European languages.

I think it is obvious they are extrapolating the traditional (not that well-known) linguistic picture of Iberia during the Iron Age, believing in continuity of that picture (especially non-Indo-European languages) during the Urnfield period and earlier.

What this data shows is, as expected, the arrival of Celtic languages in Iberia after Bell Beakers and, by extension, in the rest of western Europe. Somewhat surprisingly, this may have happened during the Urnfield period, and not during the La Tène period.

Also important are the precise subclades:

We thus detect three Bronze Age males who belonged to DF27 (154, 155), confirming its presence in Bronze Age Iberia. The other Iberian Bronze Age males could belong to DF27 as well, but the extremely low recovery rate of this SNP in our dataset prevented us to study its true distribution. All the Iberian Bronze Age males with overlapping sequences at R1b-L21 were negative for this mutation. Therefore, we can rule out Britain as a plausible proximate origin since contemporaneous British males are derived for the L21 subtype.


New open access paper Survival of Late Pleistocene Hunter-Gatherer Ancestry in the Iberian Peninsula, by Villalba-Mouco et al. Cell (2019):

BAL0051 could be assigned to haplogroup I1, while BAL003 carries the C1a1a haplogroup. To the limits of our typing resolution, EN/MN individuals CHA001, CHA003, ELT002 and ELT006 share haplogroup I2a1b, which was also reported for Loschbour [73] and Motala HG [13], and other LN and Chalcolithic individuals from Iberia [7, 9], as well as Neolithic Scotland, France, England [9], and Lithuania [14]. Both C1 and I1/ I2 are considered typical European HG lineages prior to the arrival of farming. Interestingly, CHA002 was assigned to haplogroup R1b-M343, which together with an EN individual from Cova de Els Trocs (R1b1a) confirms the presence of R1b in Western Europe prior to the expansion of steppe pastoralists that established a related male lineage in Bronze Age Europe [3, 6, 9, 13, 19]. The geographical vicinity and contemporaneity of these two sites led us to run genomic kinship analysis in order to rule out any first or second degree of relatedness. Early Neolithic individual FUC003 carries the Y haplogroup G2a2a1, commonly found in other EN males from Neolithic Anatolia [13], Starçevo, LBK Hungary [18], Impressa from Croatia and Serbia Neolithic [19] and Czech Neolithic [9], but also in MN Croatia [19] and Chalcolithic Iberia [9].

See also

The genetic and cultural barrier of the Pontic-Caspian steppe – forest-steppe ecotone

steppe-forest-steppe-biomes

We know that the Caucasus Mountains formed a persistent prehistoric barrier to cultural and population movements. Nevertheless, an even more persistent frontier to population movements in Europe, especially since the Neolithic, is the Pontic-Caspian steppe – forest-steppe ecotone.

Like the Caucasus, this barrier could certainly be crossed, and peoples and cultures could permeate in both directions, but there have been no massive migrations through it. The main connection between both regions (steppe vs. forest-steppe/forest zone) was probably through its eastern part, through the Samara region in the Middle Volga.

The chances of population expansions crossing this natural barrier anywhere else seem quite limited, with a much less porous crossing region in the west, through the Dnieper-Dniester corridor.

A Persistent ecological and cultural frontier

It is very difficult to think about any culture that transgressed this persistent ecological and cultural frontier: many prehistoric and historical steppe pastoralists did appear eventually in the neighbouring forest-steppe areas during their expansions (e.g. Yamna, Scythians, or Turks), as did forest groups who permeated to the south (e.g. Comb Ware, GAC, or Abashevo), but their respective hold in foreign biomes was mostly temporary, because their cultures had to adapt to the new ecological environment. Most if not all groups originally from a different ecological niche eventually disappeared, subjected to renewed demographic pressure from neighbouring steppe or forest populations…

The Samara region in the Middle Volga may be pointed out as the true prehistoric link between forests and steppes (see David Anthony’s remarks), something reflected in its nature as a prehistoric sink in genetics. This strong forest – forest-steppe – steppe connection was seen in the Eurasian technocomplex, during the expansion of hunter-gatherer pottery, in the expansion of Abashevo peoples to the steppes (in one of the most striking cases of population admixture in the area), with Scythians (visible in the intense contacts with Ananyino), and with Turks (Volga Turks).

steppe-forest-steppe-europe
Simplified map of the distribution of steppes and forest-steppes (Pontic and Pannonian) and xeric grasslands in Eastern Central Europe (with adjoining East European ranges) with their regionalisation as used in the review (Northern—Pannonic—Pontic). Modified from Kajtoch et al. (2016).

Before the emergence of pastoralism, the cultural contacts of the Pontic region (i.e. forest-steppes) with the Baltic were intense. In fact, the connection of the north Pontic area with the Baltic through the Dnieper-Dniester corridor and the Podolian-Volhynian region is essential to understand the spread of peoples of post-Maglemosian and post-Swiderian cultures (to the south), hunter-gatherer pottery (to the north), TRB (to the south), Late Trypillian groups (north), GAC (south), or Comb Ware (south) (see here for Eneolithic movements), and finally steppe ancestry and R1a-Z645 with Corded Ware (north). After the complex interaction of TRB, Trypillia, GAC, and CWC during the expansion of late Repin, this traditional long-range connection is lost and only emerges sporadically, such as with the expansion of East Germanic tribes.

A barrier to steppe migrations into northern Europe

One may think that this barrier was more permeable, then, in the past. However, the frontier is between steppe and forest-steppe ecological niches, and this barrier evolved during prehistory due to climate changes. The problem is, before the drought that began ca. 4000 BC and increased until the Yamna expansion, the steppe territory in the north Pontic region was much smaller, merely a strip of coastal land, compared to its greater size ca. 3300 BC and later.

This – apart from the cultural and technological changes associated with nomadic pastoralism – justifies the traditional connection of the north Pontic forest-steppes to the north, broken precisely after the expansion of Khvalynsk, as the north Pontic area became gradually a steppe region. The strips of north Pontic and Azov steppes and Crimea seem to have had stronger connections to the Northern Caucasus and Northern Caspian steppes than with the neighbouring forest-steppe areas during the Upper Palaeolithic, Mesolithic, and Neolithic.

NOTE. We still don’t know the genetic nature of Mikhailovka or Ezero, steppe-related groups possibly derived from Novodanilovka and Suvorovo close to the Black Sea (which possibly include groups from the Pannonian plains), and how they compare to neighbouring typically forest-steppe cultures of the so-called late Sredni Stog groups, like Dereivka or partly Kvityana.

steppe-forest-steppe-migration-routes
Typical migration routes through European steppes and forest-steppes. Red line represents the persistent cultural and genetic barrier, with the latest evolution in steppe region represented by the shift from dashed line to the north. Arrows show the most common population movements. Modified from Kajtoch et al. (2016).

Despite the Pontic-Caspian steppes and forest-steppes neighbouring each other for ca. 2,000 km, peoples from forested and steppe areas had an obvious advantage in their own regions, most likely due to the specialization of their subsistence economy. While this is visible already in Palaeolithic and Mesolithic hunter-gatherers, the arrival of the Neolithic package in the Pontic-Caspian region incremented the difference between groups, by spreading specialized animal domestication. The appearance of nomadic pastoralism adapted to the steppe, eventually including the use of horses and carts, made the cultural barrier based on the economic know-how even stronger.

Even though groups could still adapt and permeate a different territory (from steppe to forest-steppe/forest and vice-versa), this required an important cultural change, to the extent that it is eventually complicated to distinguish these groups from neighbouring ones (like north-west Pontic Mesolithic or Neolithic groups and their interaction with the steppes, Trypillia-Usatovo, Scythians-Thracians, etc.). In fact, this steppe – forest-steppe barrier is also seen to the east of the Urals, with the distinct expansion of Andronovo and Seima-Turbino/Andronovo-like horizons, which seem to represent completely different ethnolinguistic groups.

As a result of this cultural and genetic barrier, like that formed by the Northern Caucasus:

1) No steppe pastoralist culture (which after the emergence of Khvalynsk means almost invariably horse-riding, chariot-using nomadic herders who could easily pasture their cows in the huge grasslands without direct access to water) has ever been successful in spreading to the north or north-west into northern Europe, until the Mongols. No forest culture has ever been successful in expanding to the steppes, either (except for the infiltration of Abashevo into Sintashta-Potapovka).

2) Corded Ware was not an exception: like hunter-gatherer pottery before it (and like previous population movements of TRB, late Trypillia, GAC, Comb Ware or Lublin-Volhynia settlers) their movements between the north Pontic area and central Europe happened through forest-steppe ecological niches due to their adaptation to them. There is no reason to support a direct connection of CWC with true steppe cultures.

3) The so-called “Steppe ancestry” permeated the steppe – forest-steppe ecotone for hundreds of years during the 5th and early 4th millennium BC, due to the complex interaction of different groups, and probably to the aridization trend that expanded steppe (and probably forest-steppe) to the north. Language, culture, and paternal lineages did not cross that frontier, though.

EDIT (4 FEB 2019): Wang et al. is out in Nature Communications. They deleted the Yamna Hungary samples and related analyses, but it’s interesting to see where exactly they think the trajectory of admixture of Yamna with European MN cultures fits best. This path could also be inferred long ago from the steppe connections shown by the Yamna Hungary -> Bell Beaker evolution and by early Balkan samples:

wang-yamna-connection
Prehistoric individuals projected onto a PCA of 84 modern-day West Eurasian populations (open symbols). Dashed arrows indicate trajectories of admixture: EHG—CHG (petrol), Yamnaya—Central European MN (pink), Steppe—Caucasus (green), and Iran Neolithic—Anatolian Neolithic (brown). Modified from the original, a red circle has been added to the Yamna-Central European MN admixture.

Related

Eastern pressure blade technology in west Scandinavia associated with WHG

New interesting preprint Ancient DNA from chewing gums connects material culture and genetics of Mesolithic hunter-gatherers in Scandinavia, by Kashuba et al. (2018).

Interesting excerpts (emphasis mine):

Mitochondrial genomes from all three individuals belong to the U5a2d haplogroup. (…) The mitochondrial U5a2d haplogroup is consistent with earlier published results for ancient individuals from Scandinavia, U5a being the most common within SHG. Of the 16 Mesolithic individuals from Scandinavia published prior to our study, seven belong to the U5a haplogroup, nine share the U2 and U4 haplogroups

We divided the SHG group into two groups: SHGa and SHGb (ancient individuals found in contemporary Norway and Sweden, respectively). We based this on both the geographical distribution and the previous studies demonstrating the close relation of SHGa to EHG group and SHGb to WHG group. To further explore the demography within the SHG group, we compared the ancestry of BLE individuals within SHGa and SHGb groups. This comparison revealed a high relative shared drift between BLE individuals and the SHGb group

scandinavia-hunter-gatherer-admixture
Admixture analysis showing the major mode for K=15. The figure represents 11 runs out of 20 replicates (Greedy algorithm ran with the Jaccard distance and a 0.97 similarity threshold)

The results from Huseby Kiev allow us to finally connect the SHG group with the eastern pressure blade technology. However, the higher genetic affinity between Huseby Kiev individuals and the WHG group challenges the earlier suggested tie between eastern technology and EHG genetics. Our results suggest either early cultural transmission, or a more complex course of events involving both non- and co-dependent cultural and genetic admixture.

huseby-kiev

Seeing how culture is indeed usually associated with the expansion of a certain population, especially at such an early date, I guess this similarity with WHG of incoming eastern peoples comes from an originally EHG population expanding into a mainly WHG area in the west (similar to what happens e.g. with Bell Beakers), or being replaced later by a WHG population which adopted the culture (similar to what happened with late Corded Ware populations in central-east Europe after the expansion of Bell Beakers).

Unlike later periods, it will always be difficult to judge such ancient population movements with few samples covering thousands of years… Probably specific Y-DNA haplogroups would help differentiate between both expanding populations from east and west.

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