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 by 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

“Steppe ancestry” step by step: Khvalynsk, Sredni Stog, Repin, Yamna, Corded Ware

dzudzuana_pca-large

Wang et al. (2018) is obviously a game changer in many aspects. I have already written about the upcoming Yamna Hungary samples, about the new Steppe_Eneolithic and Caucasus Eneolithic keystones, and about the upcoming Greece Neolithic samples with steppe ancestry.

An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are in orange: equivalent Steppe_Maykop ADMIXTURE; in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups.”

Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).

1. Samara to Early Khvalynsk

The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).

Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.

PCA-caucasus-steppe-samara

This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:

steppe-maykop-admixture

NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.

2. Early Khvalynsk expansion

We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).

We also have indirect data. First, there is the PCA with outliers:

PCA-khvalynsk-steppe

Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).

Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).

3. Proto-Corded Ware expansion

It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.

Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).

Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.

NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.

PCA-sredni-stog-steppe

The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.

steppe-ancestry-admixture-sredni-stog

4. Repin / Early Yamna expansion

We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.

afanasevo-admixture

Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:

PCA-repin-yamna

This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:

yamnaya-admixture

5. Corded Ware

Corded Ware represents a quite homogeneous expansion of a late Sredni Stog population, compatible with the traditional location of Proto-Corded Ware peoples in the steppe-forest/forest zone of the Dnieper-Dniester region.

PCA-latvia-ln-steppe

We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:

sintashta-poltavka-andronovo-admixture

The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.

NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.

steppe-ancestry-admixture-latvia

A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.

NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.

Conclusion

Yamna and Corded Ware show a similar “steppe ancestry” due to convergence. I have said so many times (see e.g. here). This was clear long ago, just by looking at the Y-chromosome bottlenecks that differentiate them – and Tomenable noticed this difference in ADMIXTURE from the supplementary materials in Mathieson et al. (2017), well before Wang et al. (2018).

This different stock stems from (1) completely different ancestral populations + (2) different, long-lasting Y-chromosome bottlenecks. Their similarities come from the two neighbouring cultures admixing with similar populations.

If all this does not mean anything, and each lab was going to support some pre-selected archaeological theories from the 1960s or the 1980s, coupled with outdated linguistic models no matter what – Anthony’s model + Ringe’s glottochronological tree of the early 2000s in the Reich Lab; and worse, Kristiansen’s CWC-IE + Germano-Slavonic models of the 1940s in the Copenhagen group – , I have to repeat my question again:

What’s (so much published) ancient DNA useful for, exactly?

See also

Related

Dzudzuana, Sidelkino, and the Caucasus contribution to the Pontic-Caspian steppe

hunter-gatherer-pottery

It has been known for a long time that the Caucasus must have hosted many (at least partially) isolated populations, probably helped by geographical boundaries, setting it apart from open Eurasian areas.

David Reich writes in his book the following about India:

The genetic data told a clear story. Around a third of Indian groups experienced population bottlenecks as strong or stronger than the ones that occurred among Finns or Ashkenazi Jews. We later confirmed this finding in an even larger dataset that we collected working with Thangaraj: genetic data from more than 250 jati groups spread throughout India (…)

Rather than an invention of colonialism as Dirks suggested, long-term endogamy as embodied in India today in the institution of caste has been overwhelmingly important for millennia. (…)

The Han Chinese are truly a large population. They have been mixing freely for thousands of years. In contrast, there are few if any Indian groups that are demographically very large, and the degree of genetic differentiation among Indian jati groups living side by side in the same village is typically two to three times higher than the genetic differentiation between northern and southern Europeans. The truth is that India is composed of a large number of small populations.

There is little doubt now, based on findings spanning thousands of years, that the Mesolithic and Neolithic Caucasus hosted various very small populations, even if the ancestral components may be reduced to the few known to date (such as ANE, EHG, AME*, ENA, CHG, and other “deep” ancestral components).

NOTE. I will call the ancestral component of Dzudzuana/Anatolian hunter-gatherers Ancient Middle Easterner (AME), to give a clear idea of its likely extension during the Late Upper Palaeolithic, and to avoid using the more simplistic Dzudzuana, unless it is useful to mention these specific local samples.

dzudzuana-pca
Image modified from Lazaridis et al. (2018), including Caucasus, Don-Volga-Ural, and North Pontic Mesolithic-Neolithic populations. “Ancient West Eurasian population structure. (a) Geographical distribution of key ancient West Eurasian populations. (b) Temporal distribution of key ancient West Eurasian populations (approximate date in ky BP). (c) PCA of key ancient West Eurasians, including additional populations (shown with grey shells), in the space of outgroup f4-statistics (Methods).”

Genetic labs have a strong fixation with ancestry. I guess the use of complex statistical methods gives professionals and laymen alike the feeling of dealing with “Science”, as opposed to academic fields where you have to interpret data. I think language reveals a lot about the way people think, and the fact that ancestral components are called ‘lineages’ – while not wrong per se – is a clear symptom of the lack of interest in the true lineages: Y-DNA haplogroups.

Y-DNA bottlenecks

It has become quite clear that male-biased migrations are often the ones which can be confidently followed for actual population movements and ethnolinguistic identification, at least until the Iron Age. The frequently used Palaeolithic clusters offer a clear example of why ancestry does not represent what some people believe: They merely give a basic idea of sizeable population replacements by distant peoples.

Both concepts are important: sizeable and distant peoples. For example, during the Upper Palaeolithic in Europe there was a sizeable population replacement of the Aurignacian Goyet cluster by the Gravettian Vestonice cluster (probably from populations of far eastern Russia) coupled with the arrival of haplogroup I, although during the thousands of years that this material culture lasted, the previously expanded C1a2 lineages did not disappear, and there were probably different resurgence and admixture events.

Haplogroup I certainly expanded with the Gravettian culture to Iberia, where the Goyet ancestry did not change much – probably because of male-driven migrations -, to the extent that during the Magdalenian expansions haplogroup I expanded with an ancestry closer to Goyet, in what is called a ‘resurge’ of the Goyet cluster – even though there is a clear replacement of male lines.

The Villabruna (WHG) cluster is another good example. It probably spread with haplogroup R1b-L754, which – based on the extra ‘East Asian’ affinity of some samples and on modern samples from the Middle East – came probably from the east through a southern route, and not too long before the expansion of WHG likely from around the Black Sea, although this is still unclear. The finding of haplogroup I in samples of mostly WHG ancestry could confuse people that do not care about timing, sub-structured populations, and gene flow.

palaeolithic-expansions-reich
Image from David Reich’s Who We Are and How We Got Here. Having migrated out of Africa and the Near East, modern human pioneer populations spread throughout Eurasia (1). By at least thirty-nine thousand years ago, one group founded a lineage of European hunter-gatherers that persisted largely uninterrupted for more than twenty thousand years (2). Eventually, groups derived from an eastern branch of this founding population of European huntergatherers spread west (3), displaced previous groups, and were eventually themselves pushed out of northern Europe by the spread of glacial ice, shown at its maximum extent (top right). As the glaciers receded, western Europe was repeopled from the southwest (4) by a population that had managed to persist for tens of thousands of years and was related to an approximately thirty-five-thousand-year old individual from far western Europe. A later human migration, following the first strong warming period, had an even larger impact, with a spread from the southeast (5) that not only transformed the population of western Europe but also homogenized the populations of Europe and the Near East. At a single site—Goyet Caves in Belgium—ancient DNA from individuals spread over twenty thousand years reflects these transformations, with representatives from the Aurignacian, Gravettian, and Magdalenian periods.

NOTE. If you don’t understand why ‘clusters’ that span thousands of years don’t really matter for the many Palaeolithic population expansions that certainly happened among hunter-gatherers in Europe, just take a look at what happened with Bell Beakers expanding from Yamna into western Europe within 500 years.

If we don’t thread carefully when talking about population migrations, these terms are bound to confuse people. Just as the fixation on “steppe ancestry” – which marks the arrival in Chalcolithic Europe of peoples from the Pontic-Caspian region – has confused a lot of researchers to this day.

When I began to write about the Indo-European demic diffusion model, my concern was to find a single spot where a North-West Indo-European proto-language could have expanded from ca. 2000 BC (our most common guesstimate). Based on the 2015 papers, and in spite of their conclusions, I thought it had become clear that Corded Ware was not it, and it was rather Bell Beakers. I assumed that Uralic was spoken to the north (as was the traditional belief), and thus Corded Ware expanded from the forest zone, hence steppe ancestry would also be found there with other R1a lineages.

With the publication of Mathieson et al. (2017) and Olalde et al. (2017), I changed my mind, seeing how “steppe ancestry” did in fact appear quite late, hence it was likely to be the result of very specific population movements, probably directly from the Caucasus. Later, Mathieson published in a revision the sample from Alexandria of hg R1a-M417 (probably R1a-Z645, possibly Z93+), which further supported the idea that the migration of Corded Ware peoples started near the North Pontic forest-steppe (as I included in a the next revision).

The question remains the same I repeated recently, though: where do the extra Caucasus components (i.e. beyond EHG) of Eneolithic Ukraine/Corded Ware and Khvalynsk/Yamna come from?

Steppe ancestry: “EHG” + “CHG”?

About EHG ancestry

From Lazaridis et al. (2018):

Considering 2-way mixtures, we can model Karelia_HG as deriving 34 ± 2.8% of its ancestry from a Villabruna-related source, with the remainder mainly from ANE represented by the AfontovaGora3 (AG3) sample from Lake Baikal ~17kya.

AG3 was likely of haplogroup Q1a (as reported by YFull, see Genetiker), and probably the ANE ancestry found in Eastern Europe accompanied a Palaeolithic migration of Q1a2-M25 (formed ca. 22600 BC, TMRCA ca. 14300 BC).

NOTE. You can read more about the expansion of Q lineages during the Palaeolithic.

Combined with what we know about the Eneolithic Steppe and Caucasus populations – it is likely that ANE ancestry remained the most important component of some of the small ghost populations of the Caucasus until their emergence with the Lola culture.

pca-caucasus-dzudzuana
Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. Marked and labelled are the Balkan samples referenced in this text An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them. See the original file here. To understand the drawn potential Caucasus Mesolithic cluster, see above the PCA from Lazaridis et al. (2018).

The first sample we have now attributed to the EHG cluster is Sidelkino, from the Samara region (ca. 9300 BC), mtDNA U5a2. In Damgaard et al. (Science 2018), Yamnaya could be modelled as a CHG population related to Kotias Klde (54%) and the remaining from ANE population related to Sidelkino (>46%), with the following split events:

  1. A split event, where the CHG component of Yamnaya splits from KK1. The model inferred this time at 27 kya (though we note the larger models in Sections S2.12.4 and S2.12.5 inferred a more recent split time).
  2. A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
  3. A split event, where the ANE component of Yamnaya splits from Botai. We inferred this to occur 17 kya. Note that this is above the Sidelkino split time, so our model infers Yamnaya to be more closely related to the EHG Sidelkino, as expected.
  4. An ancestral split event between the CHG and ANE ancestral populations. This was inferred to occur around 40 kya.

Other samples classified as of the EHG cluster:

  • Popovo2 (ca. 6250 BC) of hg J1, mtDNA U4d – Po2 and Po4 from the same site (ca. 6550 BC) show continuity of mtDNA.
  • Karelia_HG, from Juzhnii Oleni Ostrov (ca. 6300 BC): I0211/UzOO40 (ca. 6300 BC) of hg J1(xJ1a), mtDNA U4a; and I0061/UzOO74 of hg R1a1(xR1a1a), mtDNA C1
  • UzOO77 and UzOO76 from Juzhnii Oleni Ostrov (ca. 5250 BC) of mtDNA R1b.
  • Samara_HG from Lebyanzhinka (ca. 5600 BC) of hg R1b1a, mtDNA U5a1d.

From the analysis of Lazaridis et al. (2018), we have some details about their admixture:

dzudzuana-admixture-sidelkino
Image modified from Lazaridis et al. (2018). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (Left) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown. (Right) ‘Speculative’ estimates. The highest number of sources (≤5) with admixture estimates within [0,1] are shown for each population. Some of the admixture proportions are not significantly different from 0 (Supplementary Information section 4).

About Anatolia_Neolithic ancestry

About the enigmatic Anatolia_Neolithic-related ancestry found in Pontic-Caspian steppe samples, this is what Wang et al. (2018) had to say:

We focused on model of mixture of proximal sources such as CHG and Anatolian Chalcolithic for all six groups of the Caucasus cluster (Eneolithic Caucasus, Maykop and Late Makyop, Maykop-Novosvobodnaya, Kura-Araxes, and Dolmen LBA), with admixture proportions on a genetic cline of 40-72% Anatolian Chalcolithic related and 28-60% CHG related (Supplementary Table 7). When we explored Romania_EN and Greece_Neolithic individuals as alternative southeast European sources (30-46% and 36-49%), the CHG proportions increased to 54-70% and 51-64%, respectively. We hypothesize that alternative models, replacing the Anatolian Chalcolithic individual with yet unsampled populations from eastern Anatolia, South Caucasus or northern Mesopotamia, would probably also provide a fit to the data from some of the tested Caucasus groups.

Also:

The first appearance of ‘Near Eastern farmer related ancestry’ in the steppe zone is evident in Steppe Maykop outliers. However, PCA results also suggest that Yamnaya and later groups of the West Eurasian steppe carry some farmer related ancestry as they are slightly shifted towards ‘European Neolithic groups’ in PC2 (Fig. 2D) compared to Eneolithic steppe. This is not the case for the preceding Eneolithic steppe individuals. The tilting cline is also confirmed by admixture f3-statistics, which provide statistically negative values for AG3 as one source and any Anatolian Neolithic related group as a second source

yamnaya-caucasus-dzudzuana
Modified image from Wang et al. (2018). In blue, Yamna-related populations. In red, Corded Ware-related populations, and two elevated Anatolia_Neolithic values in Yamna. Notice how only GAC-related admixture increases the Anatolian_N-related ancestry in the Yamna outlier from Ozero, and the late Yamna sample from Hungary, related to the homogeneous Yamna population. “Supplementary Table 14. P values of rank=3 and admixture proportions in modelling Steppe ancestry populations as a four-way admixture of distal sources EHG, CHG, Anatolian_Neolithic and WHG using 14 outgroups.Left populations: Steppe cluster, EHG, CHG, WHG, Anatolian_Neolithic. 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.”

Detailed exploration via D-statistics in the form of D(EHG, steppe group; X, Mbuti) and D(Samara_Eneolithic, steppe group; X, Mbuti) show significantly negative D values for most of the steppe groups when X is a member of the Caucasus cluster or one of the Levant/Anatolia farmer-related groups (Supplementary Figs. 5 and 6). In addition, we used f- and D-statistics to explore the shared ancestry with Anatolian Neolithic as well as the reciprocal relationship between Anatolian- and Iranian farmer-related ancestry for all groups of our two main clusters and relevant adjacent regions (Supplementary Fig. 4). Here, we observe an increase in farmer-related ancestry (both Anatolian and Iranian) in our Steppe cluster, ranging from Eneolithic steppe to later groups. In Middle/Late Bronze Age groups especially to the north and east we observe a further increase of Anatolian farmer related ancestry consistent with previous studies of the Poltavka, Andronovo, Srubnaya and Sintashta groups and reflecting a different process not especially related to events in the Caucasus.

(…) Surprisingly, we found that a minimum of four streams of ancestry is needed to explain all eleven steppe ancestry groups tested, including previously published ones (Fig. 2; Supplementary Table 12). Importantly, our results show a subtle contribution of both Anatolian farmer-related ancestry and WHG-related ancestry (Fig.4; Supplementary Tables 13 and 14), which was likely contributed through Middle and Late Neolithic farming groups from adjacent regions in the West. The discovery of a quite old AME ancestry has rendered this probably unnecessary, because this admixture from an Anatolian-like ghost population could be driven even by small populations from the Caucasus.

yamna-caucasus-cwc-anatolia-neolithic
Image modified from Wang et al. (2018). Marked are: in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus 1128 cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups (see also Supplementary Tables 10, 14 and 20).”

NOTE. For a detailed account of the possibilities regarding this differential admixture in the North Pontic area in contrast to the Don-Volga-Ural region, you can read the posts Sredni Stog, Proto-Corded Ware, and their “steppe admixture”, and Corded Ware culture origins: The Final Frontier.

While it is not yet fully clear, the increased Anatolian_Neolithic-like ancestry in Ukraine_Eneolithic samples (see below) makes it unlikely that all such ancestry in Corded Ware groups comes from a GAC-related contribution. It is likely that at least part of it represents contributions from populations of the Caucasus, based on the mostly westward population movements in the steppe from ca. 4600 BC on, including the Suvorovo-Novodanilovka expansion, and especially the Kuban-Maykop expansion during the final Eneolithic into the North Pontic area.

NOTE. Since CHG-like groups from the Caucasus may have combinations of AME and ANE ancestry similar to Yamna (which may thus appear as ‘steppe ancestry’ in the North Pontic area), it is impossible to interpret with precision the following ADMIXTURE graphic:

ukraine-whg-ehg-steppe
Modified image from Mathieson et al. (2018). Supervised ADMIXTURE analysis, modelling each ancient individual (one per row) as a mixture of population clusters constrained to contain northwestern-Anatolian Neolithic (grey), Yamnaya from Samara (yellow), EHG (pink) and WHG (green) populations. Dates in parentheses indicate approximate range of individuals in each population.

North-Eastern Technocomplex

The East Asian contribution to samples from the WHG samples (like Loschbour or La Braña), as specified in Fu et al. (2016), does not seem to be related to Baikal_EN, and appears possibly (in the ADMIXTURE analysis) integrated into he Villabruna component. I guess this implies that the shared alleles with East Asians are quite early, and potentially due to the expansion of R1b-L754 from the East.

It would be interesting to know the specific material culture Sidelkino belonged to – i.e. if it was related to the expansion of the North-Eastern Technocomplex – , and its Y-DNA. The Post-Swiderian expansion into eastern Europe, probably associated with the expansion of R1b-P297 lineages (including R1b-M73, found later in Botai and in Baltic HG) is supposed to have begun during the 11th millennium BC, but migrations to the Urals and beyond are probably concentrated in the 9th millennium, so this sample is possibly slightly early for R1b.

NOTE. User Rozenfeld at Anthrogenica posted this, which I think is interesting (in case anyone wants to try a Y-SNP call):

there is something strange with Sidelkino EHG: first, its archaeological context is not described in the supplementary. Second, its sex is not listed in the supplementary tables. Third, after looking for info about this sample, I found that: “Сиделькино-3. Для снятия вопроса о половой принадлежности индивида была проведена генетическая экспертиза, выявившая принадлежность останков мужчине.”(translation: Sidelkino-3. To resolve the question about sex of the remains, the genetic analysis was conducted, which showed that remains belonged to male), source: http://static.iea.ras.ru/books/7487_Traditsii.pdf

So either they haven’t mentioned his Y-DNA in the paper for some reason, or there are more than one Sidelkino sample and the male one has not yet been published. The coverage of the Sidelkino sample from the paper is 2.9, more than enough to tell Y-DNA haplogroup.

zaliznyak-post-swiderian
The map of spreading of Post-Swiderian and Post-Krasnosillian sites in Mesolithic of Eastern Europe in the 8th millennia BC. From Zaliznyak (see here).

My speculative guess right now about specific population movements in far eastern Europe, based on the few data we have:

  • The expansion of the North-Eastern Technocomplex first around the 9th millennium BC, most likely expanded R1b-P279 ca. 11300 BC, judging by its TMRCA, with both R1b-M73 (TMRCA 5300) and R1b-M269 (TMRCA 4400 BC) info (with extra El Mirón ancestry) back, and thus Eurasiatic.
  • The expansion of haplogroup J1 to the north may have happened before or after the R1b-P279 expansion. Judging by the increase in AG3-related ancestry near Karelia compared to Baltic_HG, it is possible that it expanded just after R1b-P279 (hence possibly J1-Y6304? TMRCA 9700 BC). Its long-lasting presence in the Caucasus is supported by the Satsurblia (ca. 11300 BC) and the Dolmen BA (ca. 1300 BC) samples.
  • The expansion of R1a-M17 ca. 6600 BC is still likely to have happened from the east, based on the R1a-M17 samples found in Baikalic cultures slightly later (ca. 5300 BC). The presence of elevated Baikal_EN ancestry in Karelia HG and in Samara HG, and the finding of R1a-M417 samples in the Forest Zone after the Mesolithic suggests a connection with the expansion of Hunter-Gatherer pottery, from the Elshanka culture in the Samara region northward into the Forset Zone and westward into the North Pontic area.
  • The expansion of R1b-M73 ca. 5300 BC is likely to be associated with the emergence of a group east of the Urals (related to the later Botai culture, and potentially Pre-Yukaghir). Its presence in a Narva sample from Donkalnis (ca. 5200 BC) suggest either an early split and spread of both R1b-P297 lineages (M73 and M269) through Eastern Europe, or maybe a back-migration with hunter-gatherer pottery.
  • R1b-M269 spread successfully ca. 4400 BC (and R1b-L23 ca. 4100 BC, both based on TMRCA), and this successful expansion is probably to be associated with the Khvalynsk-Novodanilovka expansion. We already know that Samara_HG ca. 5600 was R1b1a, so it is likely that R1b-M269 appeared (or ‘resurged’) in the Volga-Ural region shortly after the expansion of R1a-M17, whose expansion through the region may be inferred by the additional AG3 and Baikal_EN ancestry. Interesting from Samara_HG compared to the previous Sidelkino sample is the introduction of more El Mirón-related ancestry, typical of WHG populations (and thus proper of Baltic groups).

NOTE. The TMRCA dates are obviously gross approximations, because a) the actual rate of mutation is unknown and b) TMRCA estimates are based on the convergence of lineages that survived. The potential finding of R1a-Z645 (possibly Z93+) in Ukraine Eneolithic (ca. 4000 BC), and the potential finding of R1b-L23 in Khvalynsk ca. 4250 BC complicates things further, in terms of dates and origins of any subclade.

The question thus remains as it was long ago: did R1b-M269 lineages expand (‘return’) from the east, near the Urals, or directly from the north? Were they already near Samara at the same time as the expansion of hunter-gatherer pottery, and were not much affected by it? Or did they ‘resurge’ from populations admixed with Caucasus-related ancestry after the expansion of R1a-M17 with this pottery (since there are different stepped expansions from the Samara region)? We could even ask, did R1a-M17 really expand from the east, i.e. are the dates on Baikalic subclades from Moussa et al. (2016) reliable? Or did R1a-M17 expand from some pockets in the Pontic-Caspian steppe, taking over the expansion of HG pottery at some point?

hunger-gatherer-pottery
Early Neolithic cultures in eastern and central Europe: 1–Yelshanian; 2–North Caspian; 3–Rakushechnyj Yar; 4–Surskian; 5–Dnieper-Donetsian; 6– Bug-Dniesterian; 7–Upper Volga; 8–Narvian; 9–Linear Pottery. White arrows: expansion of early farming; black arrows: spread of pottery-making traditions. From Dolukhanov et al. (2009).

Maglemose-related migrations

The most interesting aspect from the new paper (regarding Indo-Uralic migrations) is that Ancestral Middle Easterner ancestry will probably be a better proxy for the Anatolia_Neolithic component found in Ukraine Mesolithic to Eneolithic, and possibly also for some of the “more CHG-like” component found among Pontic-Caspian steppe populations, all likely derived from different admixture events with groups from the Caucasus.

NOTE. Even the supposed gene flow of Neolithic Iranian ancestry into the Caucasus can be put into question, since that means possibly a Dzudzuana-like population with greater “deep ancestry” proportion than the one found in CHG, which may still be found within the Caucasus.

If it was not clear already that following ‘steppe ancestry’ wherever it appears is a rather lame way of following Indo-European migrations, every single sample from the Caucasus and their admixture with Pontic-Caspian steppe populations will probably show that “steppe ancestry” is in fact formed by a variety of steppe-related ancestral components, impossible to follow coherently with a single population. Exactly what is happening already with the Siberian ancestry.

If the paper on the Dzudzuana samples has shown something, is that the expansion of an ANE-like population shook the entire Caucasus area up to the Zagros Mountains, creating this ANE – AME cline that are CHG and Iran_N, with further contributions of “deep ancestries” (probably from the south) complicating the picture further.

If this happens with few known samples, and we know of an ANE-like ghost population in the Caucasus (appearing later in the Lola culture), we can already guess that the often repeated “CHG component” found in Ukraine_Eneolithic and Khvalynsk will not be the same (except the part mediated by the Novodanilovka expansion).

This ANE-like expansion happened probably in the Late Upper Palaeolithic, and reached Northern Europe probably after the expansion of the Villabruna cluster (ca. 12000 BC), judging by the advance of AG3-like and ENA-like ancestry in later WHG samples.

The population movements during the Mesolithic and Early Neolithic in the North Pontic area are quite complicated: the extra AME ancestry is probably connected to the admixture with populations from the Caucasus, while the close similarity of Ukraine populations with Scandinavian ones (with an increase in Villabruna ancestry from Mesolithic to Neolithic samples), probably reveal population movements related to the expansion of Maglemose-related groups.

maglemose-mesolithic
Etno-cultural situation in Central and Eastern Europe in the Late Mesolithic — Early Neolithic (VI—V Mill. BC) (after Конча 2004: 201, карта 1; made after ideas by L. L. Zaliznyak). Legend: 1 — Maglemose circle in the VII Mill. BC (after Gr. Clark); 2—7 — Mesolithic cultures of the Post-Maglemose tradition, VI Mill. BC (after S. Kozłowsky, L. L. Zaliznyak): 2 — de Leyen-Wartena; 3 — Oldesloe — Godenaa; 4 — Chojnice — Peńki; 5 — Janisłavice; 6 — finds of Janisłavice artefacts outside of the main area; 7 — Donets culture; 8 — directions of the settling of Janisłavice people (after S. Kozłowsky and L. L. Zaliznyak); 9 — the south border of Mesolithic and Early Neolithic cultures of post-Swidrian and post-Arensburgian traditions; 10 — northern border of settlement of the Balkan-Danubian farmers; 11 — Bug- Dniester culture; 12 — Neolithic cultures emerged on the ethno-cultural basis of post-Maglemose: Э — Ertebölle-Ellerbeck, Н — Neman, Д — Dnieper-Donets, М — Mariupol (western variants). From Klein (2017).

These Maglemose-related groups were probably migrants from the north-west, originally from the Northern European Plains, who occupied the previous Swiderian territory, and then expanded into the North Pontic area. The overwhelming presence of I2a (likely all I2a2a1b1b) lineages in Ukraine Neolithic supports this migration.

The likely picture of Mesolithic-Neolithic migrations in the North Pontic area right now is then:

  1. Expansion of R1a-M459 from the east ca. 12000 BC – probably coupled with AG3 and also some Baikal_EN ancestry. First sample is I1819 from Vasilievka (ca. 8700 BC), another is from Dereivka ca. 6900 BC.
  2. Expansion of R1b-V88 from the Balkans in the west ca. 9700 BC, based on its TMRCA and also the Balkan hunter-gatherer population overwhemingly of this haplogroup from the 10th millennium until the Neolithic. First sample is I1734 from Vasilievka (ca. 7252 BC), which suggests that it replaced the male population there, based on their similar EHG-like adxmixture (and lack of sizeable WHG increase), and shared mtDNA U5b2, U5a2.
  3. Expansion of I2a-Y5606 probably ca. 6800 based on its TMRCA with Janislawice culture. Supporting this is the increase in WHG contribution to Neolithic samples, including the spread of U4 subclades compared to the previous period.
  4. Expansion of R1a-M17 starting probably ca. 6600 BC in the east (see above).

NOTE. The first sample of haplogroup I appears in the Mesolithic: I1763 (ca. 8100 BC) of haplogroup I2a1, probably related to an older Upper Palaeolithic expansion.

janislawice
Distribution of archeological cultures in the North Pontic Region during the Mesolithic (7th – 6th millennium BCE). Dotted, dashed and solid lines with corresponding arrows indicate alternative models of the spread of the Grebenyky culture groups. (After Bryuako IV., Samojlova TL., Eds, Drevnie kul’tury Severo-­‐Zapadnogo Prichernomor’ya, Odessa: SMIL, 2013.) Nikitin – Ivanova 2017.

Conclusion

It is becoming more and more clear with each new paper that – unless the number of very ancient samples increases – the use of Y-chromosome haplogroups remains one of the most important tools for academics; this is especially so in the steppes, in light of the diversity found in populations from the Caucasus. A clear example comes from the Yamna – Corded Ware similarities:

After the publication of the 2015 papers, it was likely that Yamna expanded with haplogroup R1b-L23, but it has only become crystal clear that Yamna expanded through the steppes into Bell Beakers, now that we have data about the strict genetic homogeneity of the whole Yamna population from west to east (including Afanasevo), in contrast with contemporary Corded Ware peoples which expanded from a different forest-steppe population.

The presence of haplogroups Q and R1a-M459 (xM17) in Khvalynsk along with a R1b1a sample, which some interpreted as being akin to modern ‘mixed’ populations in the past, is likely to point instead to a period of Khvalynsk-Novodanilovka expansion with R1b-M269, where different small populations from the steppe were being integrated into the common Khvalynsk stock, but where differences are seen in material culture surrounding their burials, as supported by the finding of R1b1 in the Kuban area already in the first half of the 5th millennium. The case would be similar to the early ‘mixed’ Icelandic population.

Only after the emergence of the Samara culture (in the second half of the 6th millennium BC), with a sample of haplogroup R1b1a, starts then the obvious connection with Early Proto-Indo-Europeans; and only after the appearance of late Sredni Stog and haplogroup R1a-M417 (ca. 4000 BC) is its connection with Uralic also clear. In previous population movements, I think more haplogroups were involved in migrations of small groups, and only some communities among them were eventually successful, expanding to be dominant, creating ever growing cultures during their expansions.

Indeed, if you think in terms of Uralic and Indo-European just as converging languages, and forget their potential genetic connection, then the genetic + linguistic picture becomes simplified, and the upper frontier of the 6th millennium BC with a division North Pontic (Mariupol) vs. Volga-Ural (Samara) is enough. However, tracing their movements backwards – with cultural expansions from west to east (with the expansion of farming), and earlier east to west (with hunter-gatherer pottery), and still earlier west to east (with the north-eastern technocomplex), offers an interesting way to prove their potential connection to macrofamilies, at least in terms of population movements.

corded-ware-uralic-qpgraph
Modified image from Tambets et al. (2018) Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. a The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. The NeolL (Neolithic Levant) ancestry selected in this qpGraph is likely to correspond (at least in part) to a specific Dzudzuana-like component present in the CHG-like population that admixed in the North Pontic area.

I am quite convinced right now that it would be possible to connect the expansion of R1b-L754 subclades with a speculative Nostratic (given the R1b-V88 connection with Afroasiatic, and the obvious connection of R1b-L297 with Eurasiatic). Paradoxically, the connection of an Indo-Uralic community in the steppes (after the separation of Yukaghir) with any lineage expansion (R1a-M17, R1b-M269, or even Q, I or J1) seems somehow blurrier than one year ago, possibly just because there are too many open possibilities.

David Reich says about the admixture with Neanderthals, which he helped discover:

At the conclusion of the Neanderthal genome project, I am still amazed by the surprises we encountered. Having found the first evidence of interbreeding between Neanderthals and modern humans, I continue to have nightmares that the finding is some kind of mistake. But the data are sternly consistent: the evidence for Neanderthal interbreeding turns out to be everywhere. As we continue to do genetic work, we keep encountering more and more patterns that reflect the extraordinary impact this interbreeding has had on the genomes of people living today.

I think this is a shared feeling among many of us who have made proposals about anything, to fear that we have made a gross, evident mistake, and constantly look for flaws. However, it seems to me that geneticists are more preoccupied with being wrong in their developed statistical methods, in the theoretical models they are creating, and not so much about errors in the true ancient ethnolinguistic picture human population genetics is (at least in theory) concerned about. Their publications are, after all, constantly associating genetic finds with cultures and (whenever possible) languages, so this aspect of their research should not be taken lightly.

Seeing how David Anthony or Razib Khan (among many others) have changed their previously preferred migration models as new data was published, and they continue to be respected in their own fields, I guess we can be confident that professionals with integrity are going to accept whatever new picture appears. While I don’t think that genetic finds can change what we can reconstruct with comparative grammar, I am also ready to revise guesstimates and routes of expansion of certain dialects if R1a-Z645 is shown to have accompanied Late Proto-Indo-Europeans during their expansion with Yamna, and later integrated somehow with Corded Ware.

However, taking into account the obsession of some with an ancestral, uninterrupted R1a—Indo-European association, and the lack of actual political repercussion of Neanderthal admixture, I think the most common nightmare that all genetic researchers should be worried about is to keep inflating this “Yamnaya ancestry”-based hornet’s nest, which has been constantly stirred up for the past two years, by rejecting it – or, rather, specifying it into its true complex nature.

This succession of corrections and redefinitions, coupled with the distinct Y-DNA bottleneck of each steppe population, will eventually lead to a completely different ethnolinguistic picture of the Pontic-Caspian region during the Eneolithic, which is likely to eventually piss off not only reasonable academics stubbornly attached to the CWC-IE idea, but also a part of those interested in daydreaming about their patrilineal ancestors.

Sometimes it’s better to just rip off the band-aid once and for all…

Featured image from The oldest pottery in hunter-gatherer communitiesand models of Neolithisation of Eastern Europe (2015), by Andrey Mazurkevich and Ekaterina Dolbunova.

Related

Interesting is today’s post in Ancient DNA Era: Is Male-driven Genetic Replacement always meaning Language-shift?

R1a-Z280 lineages in Srubna; and first Palaeo-Balkan R1b-Z2103?

herodotus-world-map

Scythian samples from the North Pontic area are far more complex than what could be seen at first glance. From the new Y-SNP calls we have now thanks to the publications at Molgen (see the spreadsheet) and in Anthrogenica threads, I think this is the basis to work with:

NOTE. I understand that writing a paper requires a lot of work, and probably statistical methods are the main interest of authors, editors, and reviewers. But it is difficult to comprehend how any user of open source tools can instantly offer a more complex assessment of the samples’ Y-SNP calls than professionals working on these samples for months. I think that, by now, it should be clear to everyone that Y-DNA is often as important (sometimes even more) than statistical tools to infer certain population movements, since admixture can change within few generations of male-biased migrations, whereas haplogroups can’t…

Srubna

Srubna-Andronovo samples are as homogeneous as they always were, dominated by R1a-Z645 subclades and CWC-related (steppe_MLBA) ancestry.

The appearance of one (possibly two) R-Z280 lineages in this mixed Srubna-Alakul region of the southern Urals and this early (1880-1690 BC, hence rather Pokrovka-Alakul) points to the admixture of R1a-Z93 and R1a-Z280 already in Abashevo, which also explains the wide distribution of both subclades in the forest zones of Central Asia.

If Abashevo is the cornerstone of the Indo-Iranian / Uralic community, as it seems, the genetic admixture would initially be quite similar, undergoing in the steppes a reduction to haplogroup R1a-Z93 (obviously not complete), at the same time as it expanded to the west with Pokrovka and Srubna, and to the east with Petrovka and Andronovo. To the north, similar reductions will probably be seen following the Seima-Turbino phenomenon.

NOTE. Another R1a-Z280 has been found in the recent sample from Bronze Age Poland (see spreadsheet). As it appears right now in ancient and modern DNA, there seems to be a different distribution between subclades:

  • R1a-Z280 (formed ca. 2900 BC, TMRCA ca. 2600 BC) appears mainly distributed today to the east, in the forest and steppe regions, with the most ‘successful’ expansions possibly related to the spread of Abashevo- and Battle Axe-related cultures (Indo-Iranian and Uralic alike).
  • R1a-M458 (formed ca. 2700, TMRCA ca. 2700 BC) appears mainly distributed to the north, from central Europe to the east – but not in the steppe in aDNA, with the most ‘successful’ expansions to the west.

M458 lineages seem thus to have expanded in the steppe in sizeable numbers only after the Iranian expansions (see a map of modern R1a distributions) i.e. possibly with the expansion of Slavs, which supports the model whereby cultures from central-east Europe (like Trzciniec and Lusatian), accompanied mainly by M458 lineages, were responsible for the expansion of Proto-Balto-Slavic (and later Proto-Slavic).

The finding of haplogroup R1a-Z93, among them one Z2123, is no surprise at this point after other similar Srubna samples. As I said, the early Srubna expansion is most likely responsible for the Szólád Bronze Age sample (ca. 2100-1700 BC), and for the Balkans BA sample (ca. 1750-1625 BC) from Merichleri, due to incursions along the central-east European steppe.

cheek-pieces
Map of decorated bone/antler bridle cheek-pieces and whip handle equivalents. They are often local translations that remained faithful to the originals (from data in Piggott, 1965; Kristiansen & Larsson, 2005; David, 2007). Image from Vandkilde (2014).

Cimmerians

Cimmerian samples from the west show signs of continuity with R1a-Z93 lineages. Nevertheless, the sample of haplogroup Q1a-Y558, together with the ‘Pre-Scythian’ sample of haplogroup N (of the Mezőcsát Culture) in Hungary ca. 980-830 BC, as well as their PCA, seem to depict an origin of these Pre-Scythian peoples in populations related to the eastern Central Asian steppes, too.

NOTE. I will write more on different movements (unrelated to Uralic expansions) from Central and East Asia to the west accompanied by Siberian ancestry and haplogroup N with the post of Ugric-Samoyedic expansions.

Scythians

The Scythian of Z2123 lineage ca. 375-203 BC from the Volga (in Mathieson et al. 2015), together with the sample scy193 from Glinoe (probably also R1a-Z2123), without a date, as well as their common Steppe_MLBA cluster, suggest that Scythians, too, were at first probably quite homogeneous as is common among pastoralist nomads, and came thus from the Central Asian steppes.

The reduction in haplogroup variability among East Iranian peoples seems supported by the three new Late Sarmatian samples of haplogroup R1a-Z2124.

Approximate location of Glinoe and Glinoe Sad (with Starosilya to the south, in Ukrainian territory):

This initial expansion of Scythians does not mean that one can dismiss the western samples as non-Scythians, though, because ‘Scythian’ is a cultural attribution, based on materials. Confirming the diversity among western Scythians, a session at the recent ISBA 8:

Genetic continuity in the western Eurasian Steppe broken not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths), by Järve et al.

The long-held archaeological view sees the Early Iron Age nomadic Scythians expanding west from their Altai region homeland across the Eurasian Steppe until they reached the Ponto-Caspian region north of the Black and Caspian Seas by around 2,900 BP. However, the migration theory has not found support from ancient DNA evidence, and it is still unclear how much of the Scythian dominance in the Eurasian Steppe was due to movements of people and how much reflected cultural diffusion and elite dominance. We present new whole-genome results of 31 ancient Western and Eastern Scythians as well as samples pre- and postdating them that allow us to set the Scythians in a temporal context by comparing the Western Scythians to samples before and after within the Ponto-Caspian region. We detect no significant contribution of the Scythians to the Early Iron Age Ponto-Caspian gene pool, inferring instead a genetic continuity in the western Eurasian Steppe that persisted from at least 4,800–4,400 cal BP to 2,700–2,100 cal BP (based on our radiocarbon dated samples), i.e. from the Yamnaya through the Scythian period.

(…) Our results (…) support the hypothesis that the Scythian dominance was cultural rather than achieved through population replacement.

Detail of the slide with admixture of Scythian groups in Ukraine:

scythians-admixture

The findings of those 31 samples seem to support what Krzewińska et al. (2018) found in a tiny region of Moldavia-south-western Ukraine (Glinoi, Glinoi Sad, and Starosilya).

The question, then, is as follows: if Scythian dominance was “cultural rather than achieved through population replacement”…Where are the R1b-Z2103 from? One possibility, as I said in the previous post, is that they represent pockets of Iranian R1b lineages in the steppes descended from eastern Yamna, given that this haplogroup appears in modern populations from a wide region surrounding the steppes.

The other possibility, which is what some have proposed since the publication of the paper, is that they are related to Thracians, and thus to Palaeo-Balkan populations. About the previously published Thracian individuals in Sikora et al. (2014):

thracian-samples
Geographic origin of ancient samples and ADMIXTURE results. (A) Map of Europe indicating the discovery sites for each of the ancient samples used in this study. (B) Ancestral population clusters inferred using ADMIXTURE on the HGDP dataset, for k = 6 ancestral clusters. The width of the bars of the ancient samples was increased to aid visualization. https://doi.org/10.1371/journal.pgen.1004353.g001

For the Thracian individuals from Bulgaria, no clear pattern emerges. While P192-1 still shows the highest proportion of Sardinian ancestry, K8 more resembles the HG individuals, with a high fraction of Russian ancestry.

Despite their different geographic origins, both the Swedish farmer gok4 and the Thracian P192-1 closely resemble the Iceman in their relationship with Sardinians, making it unlikely that all three individuals were recent migrants from Sardinia. Furthermore, P192-1 is an Iron Age individual from well after the arrival of the first farmers in Southeastern Europe (more than 2,000 years after the Iceman and gok4), perhaps indicating genetic continuity with the early farmers in this region. The only non-HG individual not following this pattern is K8 from Bulgaria. Interestingly, this individual was excavated from an aristocratic inhumation burial containing rich grave goods, indicating a high social standing, as opposed to the other individual, who was found in a pit.

pca-thracians

The following are excerpts from A Companion to Ancient Thrace (2015), by Valeva, Nankov, and Graninger (emphasis mine):

Thracian settlements from the 6th c. BC on:

(…) urban centers were established in northeastern Thrace, whose development was linked to the growth of road and communication networks along with related economic and distributive functions. The early establishment of markets/emporia along the Danube took place toward the middle of the first millennium BCE (Irimia 2006, 250–253; Stoyanov in press). The abundant data for intensive trade discovered at the Getic village in Satu Nou on the right bank of the Danube provides another example of an emporion that developed along the main artery of communication toward the interior of Thrace (Conovici 2000, 75–76).

Undoubtedly the most prominent manifestation of centralization processes and stratification in the settlement system of Thrace arrives with the emergence of political capitals – the leading urban centers of various Thracian political formations.

getic-thracian
Image from Volf at Vol_Vlad LiveJournal.

Their relationships with Scythians and Greeks

The Scythian presence south of the Danube must be balanced with a Thracian presence north of the river. We have observed Getae there in Alexander’s day, settled and raising grain. For Strabo the coastlands from the Danube delta north as far as the river and Greek city of Tyras were the Desert of the Getae (7.3.14), notable for its poverty and tracklessness beyond the great river. He seems to suggest also that it was here that Lysimachus was taken alive by Dromichaetes, king of the Getae, whose famous homily on poverty and imperialism only makes sense on the steppe beyond the river (7.3.8; cf. Diod. 21.12; further on Getic possessions above the Danube, Paus. 1.9 with Delev 2000, 393, who seems rather too skeptical; on poverty, cf. Ballesteros Pastor 2003). This was the kind of discourse more familiarly found among Scythians, proud and blunt in the strength of their poverty. However, as Herodotus makes clear, simple pastoralism was not the whole story as one advanced round into Scythia. For he observes the agriculture practiced north and west of Olbia. These were the lands of the Alizones and the people he calls the Scythian Ploughmen, not least to distinguish them from the Royal Scythians east of Olbia, in whose outlook, he says, these agriculturalist Scythians were their inferiors, their slaves (Hdt. 4.20). The key point here is that, as we began to see with the Getan grain-fields of Alexander’s day, there was scope for Thracian agriculturalists to maintain their lifestyles if they moved north of the Danube, the steppe notwithstanding. It is true that it is movement in the other direction that tends to catch the eye, but there are indications in the literary tradition and, especially, in the archaeological record that there was also significant movement northward from Thrace across the Danube and the Desert of the Getae beyond it.

Greek literary sources were not much concerned with Thracian migration into Scythia, but we should observe the occasional indications of that process in very different texts and contexts. At the level of myth, it is to be remembered that Amazons were regularly considered to be of Thracian ethnicity from Archaic times onward and so are often depicted in Thracian dress in Greek art (Bothmer 1957; cf. Sparkes 1997): while they are most familiar on the south coast of the Black Sea, east of Sinope, they were also located on the north coast, especially east of the Don (the ancient Tanais). Herodotus reports an origin-story of the Sauromatians there, according to which this people had been created by the union of some Scythian warriors with Amazons captured on the south coast and then washed up on the coast of Scythia (4.110). While the story is unhistorical, it is not without importance. First, it reminds us that passage north from the Danube was not the only way that Thracians, Thracian influence, and Thracian culture might find their way into Scythia. There were many more and less circuitous routes, especially by sea, that could bring Thrace into Scythia. Secondly, the myth offered some ideological basis for the Sauromatian settlement in Thrace that Strabo records, for Sauromatians might claim a Thracian origin through their Amazon forebears. Finally, rather as we saw that Heracles could bring together some of the peoples of the region, we should also observe that Ares, whose earthly home was located in Thrace by a strong Greek and Roman tradition, seems also to have been a deity of special significance and special cult among the Scythians. So much was appropriate, especially from a Classical perspective, in associations between these two peoples, whose fame resided especially in their capacity for war.

skythen
Scythians: cultures and findings (ca. 7th-4th/3rd c. BC). Greek colonies marked with concentric circles.

This broad picture of cultural contact, interaction, and osmosis, beyond simple conflict, provides the context for a range of archaeological discoveries, which – if examined separately – may seem to offer no more than a scatter of peculiarities. Here we must acknowledge especially the pioneering work of Melyukova, who has done most to develop thinking on Thracian–Scythian interaction. As she pointed out, we have a good example of Thracian–Scythian osmosis as early as the mid-seventh century bce at Tsarev Brod in northeastern Bulgaria, where a warrior’s burial combines elements of Scythian and Thracian culture (Melyukova 1965). For, while the manner of his burial and many of the grave goods find parallels in Scythia and not Thrace, there are also goods which would be odd in a Scythian burial and more at home in a Thracian one of this period (notably a Hallstatt vessel, an iron knife, and a gold diadem). Also interesting in this regard are several stone figures found in the Dobrudja which resemble very closely figures of this kind (baby) known from Scythia (Melyukova 1965, 37–38). They range in date from perhaps the sixth to the third centuries bce, and presumably were used there – as in Scythia – to mark the burials of leading Scythians deposited in the area. Is this cultural osmosis? We should probably expect osmosis to occur in tandem with the movement of artefacts, so that only good contexts can really answer such questions from case to case. However, the broad pattern is indicated by a range of factors. Particularly notable in this regard is the observable development of a Thraco-Scythian form of what is more familiar as “Scythian animal style,” a term which – it must be understood – already embraces a range of types as we examine the different examples of the style across the great expanse from Siberia to the western Ukraine. As Melyukova observes, Thrace shows both items made in this style among Scythians and, more numerous and more interesting, a Thracian tendency to adapt that style to local tastes, with observable regional distinctions within Thrace itself. Among the Getae and Odrysians the adaptation seems to have been at its height from the later fifth century to the mid-third century (Melyukova 1965, 38; 1979).

The absence of local animal style in Bulgaria before the fifth century bce confirms that we have cultural influences and osmosis at work here, though that is not to say that Scythian tradition somehow dominated its Thracian counterpart, as has been claimed (pace Melyukova 1965, 39; contrast Kitov 1980 and 1984). Of particular interest here is the horse-gear (forehead-covers, cheek-pieces, bridle fittings, and so on) which is found extensively in Romania and Bulgaria as well as in Scythia, both in hoarded deposits and in burials. This exemplifies the development of a regional animal style, not least in silver and bronze, which problematizes the whole issue of the place(s) of its production. Accordingly, the regular designation as “Thracian” of horse-gear from the rich fourth century Scythian burial of Oguz in the Ukraine becomes at least awkward and questionable (further, Fialko 1995). And let us be clear that this is no minor matter, nor even part of a broader debate about the shared development of toreutics among Thracians and Scythians (e.g., Kitov 1980 and 1984). A finely equipped horse of fine quality was a strong statement and striking display of wealth and the power it implied

(…) while Thracian pottery appears at Olbia, Scythian pottery among Thracians is largely confined to the eastern limits of what should probably be regarded as Getic territory, namely the area close to the west of the Dniester, from the sixth century bce. Rather exceptional then is the Scythian pottery noted at Istros, which has been explained as a consequence of the Scythian pursuit of the withdrawing army of Darius and, possibly, a continued Scythian grip on the southern Danube in its aftermath (Melyukova 1965, 34). The archaeology seems to show us, therefore, that the elite Thracians and Scythians were more open to adaptation and acculturation than were their lesser brethren.

palaeo-balkan-languages
Paleo-Balkan languages in Eastern Europe between 5th and 1st century BC. From Wikipedia.

Conclusion

(…) we see distinct peoples and organizations, for example as Sitalces’ forces line up against the Scythians. Much more striking, however, against that general background, are the various ways in which the two peoples and their elites are seen to interact, connect, and share a cultural interface. We see also in Scyles’ story how the Greek cities on the coast of Thrace and Scythia played a significant role in the workings of relationships between the two peoples. It is not simply that these cities straddled the Danube, but also that they could collaborate – witness the honors for Autocles, ca. 300 bce (SEG 49.1051; Ochotnikov 2006) – and were implicated with the interactions of the much greater non-Greek powers around them. At the same time, we have seen the limited reality of familiar distinctions between settled Thracians and nomadic Scythians and the limited role of the Danube too in dividing Thrace and Scythia. The interactions of the two were not simply matters of dynastic politics and the occasional shared taste for artefacts like horse-gear, but were more profoundly rooted in the economic matrix across the region, so that “Scythian” nomadism might flourish in the Dobrudja and “Thracian-style” agriculture and settlement can be traced from Thrace across the Danube as far as Olbia. All of that offers scant justification for the Greek tendency to run together Thracians and Scythians as much the same phenomenon, not least as irrational, ferocious, and rather vulgar barbarians (e.g., Plato, Rep. 435b), because such notions were the result of ignorance and chauvinism. However, Herodotus did not share those faults to any degree, so that we may take his ready movement from Scythians to Thracians to be an indication of the importance of interaction between the two peoples whom he had encountered not only as slaves in the Aegean world, but as powerful forces in their own lands (e.g., Hdt. 4.74, where Thracian usage is suddenly brought into his account of Scythian hemp). Similarly, Thucydides, who quite without need breaks off his disquisition on the Odrysians to remark upon political disunity among the Scythians (Thuc. 2.97, a favorite theme: cf. Hdt. 4.81; Xen., Cyr. 1.1.4). As we have seen throughout this discussion, there were many reasons why Thracians might turn the thoughts of serious writers to Scythians and vice versa.

It seems, following Sikora et al. (2014), that Thracian ‘common’ populations would have more Anatolian Neolithic ancestry compared to more ‘steppe-like’ samples. But there were important differences even between the two nearby samples published from Bulgaria, which may account for the close interaction between Scythians and Thracians we see in Krzewińska et al. (2018), potentially reflected in the differences between the Central, Southern and the South-Central clusters (possibly related to different periods rather than peoples??).

If these R1b-Z2103 were descended from Thracian elites, this would be the first proof of Palaeo-Balkan populations showing mainly R1b-Z2103, as I expect. Their appearance together with haplogroup I2a2a1b1 (also found in Ukraine Neolithic and in the Yamna outlier from Bulgaria) seem to support this regional continuity, and thus a long-lasting cultural and ethnic border roughly around the Danube, similar to the one found in the northern Caucasus.

However, since these samples are some 2,500 years younger than the Yamna expansion to the south, and they are archaeologically Scythians, it is impossible to say. In any case, it would seem that the main expansion of R1a-Z645 lineages to the south of the Danube – and therefore those found among modern Greeks – was mediated by the Slavic expansions centuries later.

krzewinska-scythians-pca
Modified image from Krzewińska et al. (2018), with added Y-DNA haplogroups to each defined Scythian cluster and Sarmatians. Principal component analysis (PCA) plot visualizing 35 Bronze Age and Iron Age individuals presented in this study and in published ancient individuals in relation to modern reference panel from the Human Origins data set. See image with population references.

On the Northern cluster there is a sample of haplogroup R1b-P312 which, given its position on the PCA (apparently even more ‘modern Celtic’-like than the Hallstatt_Bylany sample from Damgaard et al. 2018), it seems that it could be the product of the previous eastward Hallstatt expansion…although potentially also from a recent one?:

Especially important in the archaeology of this interior is the large settlement at Nemirov in the wooded steppe of the western Ukraine, where there has been considerable excavation. This settlement’s origins evidently owe nothing significant to Greek influence, though the early east Greek pottery there (from ca. 650 bce onward: Vakhtina 2007) and what seems to be a Greek graffito hint at its connections with the Greeks of the coast, especially at Olbia, which lay at the estuary of the River Bug on whose middle course the site was located (Braund 2008). The main interest of the site for the present discussion, however, is its demonstrable participation in the broader Hallstatt culture to its west and south (especially Smirnova 2001). Once we consider Nemirov and the forest steppe in connection with Olbia and the other locations across the forest steppe and coastal zone, together with the less obvious movements across the steppe itself, we have a large picture of multiple connectivities in which Thrace bulks large.

scythian-peoples-balkans
Early Iron Age cultures of the Carpathian basin ca. 7-6th century BC, including steppe-related groups. Ďurkovič et al. (2018).

While the above description of clear-cut R1a-Steppe and R1b-Balkans is attractive (and probably more reliable than admixture found in scattered samples of unclear dates), the true ancient genetic picture is more complicated than that:

  • There is nothing in the material culture of the published western Scythians to distinguish the supposed Thracian elites.
  • We have the sample I0575, an Early Sarmatian from the southern Urals (one of the few available) of haplogroup R1b-Z2106, which supports the presence of R1b-Z2103 lineages among Eastern Iranian-speaking peoples.
  • We also have DA30, a Sarmatian of I2b lineage from the central steppes in Kazakhstan (ca. 47 BC – 24 AD).
  • Other Sarmatian samples of haplogroup R remain undefined.
  • There is R1a-Z93 in a late Sarmatian-Hun sample, which complicates the picture of late pastoralist nomads further.

Therefore, the possibility of hidden pockets of Iranian peoples of R1b-Z2103 (maybe also R1b-P312) lineages remains the best explanation, and should not be discarded simply because of the prevalent haplogroups among modern populations, or because of the different clusters found, or else we risk an obvious circular reasoning: “this sample is not (autosomically or in prevalent haplogroups) like those we already had from the steppe, ergo it is not from this or that steppe culture.” Hopefully, the upcoming paper by Järve et al. will help develop a clearer genetic transect of Iranian populations from the steppes.

All in all, the diversity among western Scythians represents probably one of the earliest difficult cases of acculturation to be studied with ancient DNA (obviously not the only one), since Scythians combine unclear archaeological data with limited and conflicting proto-historical accounts (also difficult to contrast with the wide confidence intervals of radiocarbon dates) with different evolving clusters and haplogroups – especially in border regions with strong and continued interactions of cultures and peoples.

With emerging complex cases like these during the Iron Age, I am happy to see that at least earlier expansions show clearer Y-DNA bottlenecks, or else genetics would only add more data to argue about potential cultural diffusion events, instead of solving questions about proto-language expansions once and for all…

Related

Y-chromosome mixture in the modern Corsican population shows different migration layers

mesolithic-europe

Open access Prehistoric migrations through the Mediterranean basin shaped Corsican Y-chromosome diversity, by Di Cristofaro et al. PLOS One (2018).

Interesting excerpts:

This study included 321 samples from men throughout Corsica; samples from Provence and Tuscany were added to the cohort. All samples were typed for 92 Y-SNPs, and Y-STRs were also analyzed.

Haplogroup R represented approximately half of the lineages in both Corsican and Tuscan samples (respectively 51.8% and 45.3%) whereas it reached 90% in Provence. Sub-clade R1b1a1a2a1a2b-U152 predominated in North Corsica whereas R1b1a1a2a1a1-U106 was present in South Corsica. Both SNPs display clinal distributions of frequency variation in Europe, the U152 branch being most frequent in Switzerland, Italy, France and Western Poland. Calibrated branch lengths from whole Y chromosome sequencing [44,45] and ancient DNA studies [46] both indicated that R1a and R1b diversification began relatively recently, about 5 Kya, consistent with Bronze Age and Copper Age demographic expansion. TMRCA estimations are concordant with such expansion in Corsica.

corsica-haplogroups
Spatial frequency maps for haplogroups with frequencies above 3%, their Y-STR based phylogenetic networks in Corsican populations (Blue: North, Green: West, Orange: South, Black: Center and Purple: East) and their TMRCA (in years, +/- SE).

Haplogroup G reached 21.7% in Corsica and 13.3% in Tuscany. Sub-clade G2a2a1a2-L91 accounted for 11.3% of all haplogroups in Corsica yet was not present in Provence or in Tuscany. Thirty-four out of the 37 G2a2a1a2-L91 displayed a unique Y-STR profile, illustrated by the star-like profile of STR networks (Fig 1). G2a2a1a2-L91 and G2a2a-PF3147(xL91xM286) show their highest frequency in present day Sardinia and southern Corsica compared to low levels from Caucasus to Southern Europe, encompassing the Near and Middle East [21,47–50]. Ancient DNA results from Early and Middle Neolithic samples reported the presence of haplogroup G2a-P15 [51–53], consistent with gene flow from the Mediterranean region during the Neolithic transition. Td expansion time estimated by STR for P15-affiliated chromosomes was estimated to be 15,082+/-2217 years ago [49]. Ötzi, the 5,300-year-old Alpine mummy, was derived for the L91 SNP [21]. A genetic relationship between G haplogroups from Corsica and Sardinia is further supported by DYS19 duplication, reported in North Sardinia [14], and observed in the southern part of the Corsica in 9 out of 37 G2a2a1a2-L91 chromosomes and in 4 out of 5 G2a2a-PF3147(xL91xM286) chromosomes, 3 of which displayed an identical STR profile (S4 Table).

This lineage has a reported coalescent age estimated by whole sequencing in Sardinian samples of about 9,000 years ago. This could reflect common ancestors coming from the Caucasus and moving westward during the Neolithic period [48], whereas their continental counterparts would have been replaced by rapidly expanding populations associated with the Bronze Age [46,54,55]. Estimated TMRCA for L91 lineage in Corsica is 4529 +/- 853 years. G-L497 showed high frequencies in Corsica compared to Provence and Tuscany, and this haplogroup was common in Europe, but rare in Greece, Anatolia and the Middle East. Fifteen out of the 17 Corsican G2a2b2a1a1b-L497 displayed a unique Y-STR profile (S4 Table) with an estimated TMRCA of 6867 +/- 1294 years. Haplogroup G2a2b1-M406, associated with Impressed Ware Neolithic markers, along with J2a1-DYS445 = 6 and J2a1b1-M92 [22,49], had very low levels in Corsica. Conversely, G2a2b2a-P303was highly represented and seemed to be independent of the G2a2b1-M406 marker. The 7 G2a2b2a-P303(xL497xM527) Corsican chromosomes displayed a unique Y-STR profile (S4 Table).

pca-corsica
First and second axes of the PCA based on 12 Y-chromosome haplogroup frequencies in 83 west Mediterranean populations.

Haplogroup J, mainly represented by J2a1b-M67(xM92), displayed intermediate frequencies in Corsica compared to Tuscany and Provence. J2a1b-M67(xM92) derived STR network analysis displayed a quite homogeneous profile across the island with an estimated TMRCA of 2381 +/- 449 years (Fig 1) and individuals displaying M67 were peripheral compared to Northwestern Italians (S2 Fig). The haplogroup J2a1-Page55(xM67xM530), characteristic of non-Greek Anatolia [22], was found in the north-west of Corsica. Haplogroup J2a1-DYS445 = 6 was found in the north-west with DYS391 = 10 repeats, and in the far south with DYS391 = 9 repeats, the former was associated with Anatolian Greek samples, whereas the second was found in central Anatolia [22]. The 7 J2b2a-M241 displayed a unique Y-STR profile (S4 Table), they were only detected in the Cap Corse region, this sub-haplogroup shows frequency peaks in both the southern Balkans and northern-central Italy [56] and is associated with expansion from the Near East to the Balkans during Neolithic period [57].

Haplogroup E, mainly represented by E1b1b1a1b1a-V13, displayed intermediate frequencies in Corsica compared to Tuscany and Provence. E1b1b1a1b1a-V13 was thought to have initiated a pan-Mediterranean expansion 7,000 years ago starting from the Balkans [52] and its dispersal to the northern shore of the Mediterranean basin is consistent with the Greek Anatolian expansion to the western Mediterranean [22], characteristic of the region surrounding Alaria, and consistent with the TMRCA estimated in Corsica for this haplogroup. A few E1b1a-V38 chromosomes are also observed in the same regions as V13.

Related:

Haplogroup J spread in the Mediterranean due to Phoenician and Greek colonizations

iron_age_europe_mediterranean

Open access A finely resolved phylogeny of Y chromosome Hg J illuminates the processes of Phoenician and Greek colonizations in the Mediterranean, by Finocchio et al. Scientific Reports (2018) Nº 7465.

Abstract (emphasis mine):

In order to improve the phylogeography of the male-specific genetic traces of Greek and Phoenician colonizations on the Northern coasts of the Mediterranean, we performed a geographically structured sampling of seven subclades of haplogroup J in Turkey, Greece and Italy. We resequenced 4.4 Mb of Y-chromosome in 58 subjects, obtaining 1079 high quality variants. We did not find a preferential coalescence of Turkish samples to ancestral nodes, contradicting the simplistic idea of a dispersal and radiation of Hg J as a whole from the Middle East. Upon calibration with an ancient Hg J chromosome, we confirmed that signs of Holocenic Hg J radiations are subtle and date mainly to the Bronze Age. We pinpointed seven variants which could potentially unveil star clusters of sequences, indicative of local expansions. By directly genotyping these variants in Hg J carriers and complementing with published resequenced chromosomes (893 subjects), we provide strong temporal and distributional evidence for markers of the Greek settlement of Magna Graecia (J2a-L397) and Phoenician migrations (rs760148062). Our work generated a minimal but robust list of evolutionarily stable markers to elucidate the demographic dynamics and spatial domains of male-mediated movements across and around the Mediterranean, in the last 6,000 years.

greek-phoenician
J2-L397. The star indicates the centroid of derived alleles. The solid square indicates the centroid of ancestral alleles, with its 95% C.I. (ellipse). In the insets: distributions of the pairwise sampling distances (in Km) for the carriers of the ancestral (black) and derived (white) allele, with solid and dashed lines indicating the respective averages. At right: median joining network of 7-STR haplotypes and SNPs in the same groups, with sectors coloured according to sampling location. Haplotype structure is detailed for some nodes, in the order YCA2a-YCA2b-DYS19-DYS390-DYS391-DYS392-DYS393 (in italics).

Interesting excerpts:

Two features of our tree are at odds with the simplistic idea of a dispersal of Hg J as a whole from the Middle East towards Greece and Italy and an accompanying radiation26. First, there is little evidence of sudden diversification between 15 and 5 kya, a period of likely population increase and pressure for range expansion, due to the Agricultural revolution in the Fertile Crescent. Second, within each subclade, lineages currently sampled in Turkey do not show up as preferentially ancestral. Both findings are replicated and reinforced by examining the previous landmark studies. Our Turkish samples do not coalesce preferentially to ancestral nodes when mapped onto these studies’ trees.

Additional relevant information on the entire Hg J comes from the discontinuous distribution of J2b-M12. The northern fringe of our sample is enriched in the J2b-M241 subclade, which reappears in the gulf of Bengal38,45, with low frequencies in the intervening Iraq46 and Iran47. No J2b-M12 carriers were found among 35 modern Lebanese, as contrasted to one of two ancient specimens from the same region35.

In summary, a first conclusion of our sequencing effort and merge with available data is that the phylogeography of Hg J is complex and hardly explained by the presence of a single population harbouring the major lineages at the onset of agriculture and spreading westward. A unifying explanation for all the above inconsistencies could be a centre of initial radiation outside the area here sampled more densely, i.e. the Caucasus and regions North of it, from which different Hg J subclades may have later reached mainland Italy, Greece and Turkey, possibly following different routes and times. Evidence in this direction comes from the distribution of J2a-M41045,48 and the early-49 or mid-Holocene50 southward spread of J1.

greek-colonization
Supplemental Figure 7. Maps of sampling locations for the carriers of the derived allele (white triangle point down) at the indicated SNP vs carriers of the ancestral allele (black triangle point-up), conditioned on identical genotype at the same most terminal marker. Coastlines were drawn with the R packages18 “map” and “mapproj” v. 3.1.3 (https://cran.r-project.org/web/packages/mapproj/index.html), and additional features added with default functions. The star triangle indicates the centroid of derived alleles. The solid square indicates the centroid of ancestral alleles, with its 95% C.I. (ellipse). In the insets: distributions of the pairwise sampling distances (in Km) for the carriers of the ancestral (black) and derived (white) allele, with solid and dashed lines indicating the respective averages. At right: median joining network of 7-STR haplotypes and SNPs in the same groups, with sectors coloured according to sampling location. Haplotype structure is detailed for some nodes, in the order YCA2a-YCA2b-DYS19-DYS390-DYS391-DYS392-DYS393 (in italics).

The lineage defined by rs779180992, belonging to J2b-M205, and dated at 4–4.5 kya, has a radically different distribution, with derived alleles in Continental Italy, Greece and Northern Turkey, and two instances in a Palestinian and a Jew. The interpretation of the spread of this lineage is not straightforward. Tentative hypotheses are linked to Southward movements that occurred in the Balkan Peninsula from the Bronze Age29,53, through the Roman occupation and later54.

The slightly older (5.6–6.3 kya) branch 98 lineage displays a similar trend of a Eastward positioning of derived alleles, with the notable difference of being present in Sardinia, Crete, Cyprus and Northern Egypt. This feature and the low frequency of the parental J2a-M92 lineage in the Balkans27 calls for an explanation different from the above.

Finally, we explored the distribution of J2a-L397 and three derived lineages within it. J2a-L397 is tightly associated with a typical DYS445 6-repeat allele. This has been hypothesized as a marker of the Greek colonizations in the Mediterranean55, based on its presence in Greek Anatolia and Provence (France), a region with attested Iron Age Greek contribution. All of our chromosomes in this clade were characterized also by DYS391(9), confirming their Anatolian Greek signature. We resolved the J2a-L397 clade to an unprecedented precision, with three internal markers which allow a finer discrimination than STRs. The ages of the three lineages (2.0–3.0 kya) are compatible with the beginning of the Greek colonial period, in the 8th century BCE. The three subclades have different distributions (Fig. 2B), with two (branches 57, 59) found both East and West to Greece, and one only in Italy (branch 58). As to Mediterranean Islands, J2a-L397 was found in Cyprus56 and Crete43. Its presence as one of the three branches 57–59 will represent an important test. In Italy all three variants were found mainly along the Western coast (18/25), which hosted the preferred Greek trade cities. The finding of all three differentiated lineages in Locri excludes a local founder effect of a single genealogy. Interestingly, an important Greek colony was established in this location, with continuity of human settlement until modern times. The sample composed of the same subjects displayed genetic affinities with Eastern Greece and the Aegean also at autosomal markers57. In summary, the distributions of branches 57–59 mirror the variety of the cities of origin and geographic ranges during the phases of the colonization process58.

So, there you have it, another proof that haplogroup J and CHG-related ancestry in the Mediterranean was mainly driven by different (and late) expansions of historic peoples.

Related:

Consequences of Damgaard et al. 2018 (I): EHG ancestry in Maykop samples, and the potential Anatolian expansion routes

neolithic_steppe-anatolian-migrations

This is part I of two posts on the most recent data concerning the earliest known Indo-European migrations.

Anatolian in Armi

I am reading in forums about “Kroonen’s proposal” of Anatolian in the 3rd millennium. That is false. The Copenhagen group (in particular the authors of the linguistic supplement, Kroonen, Barjamovic, and Peyrot) are merely referencing Archi (2011. “In Search of Armi”. Journal of Cuneiform Studies 63: 5–34) in turn using transcriptions from Bonechi (1990. “Aleppo in età arcaica; a proposito di un’opera recente”. Studi Epigrafici e Linguistici sul Vicino Oriente Antico 7: 15–37.), who asserted the potential Anatolian origin of the terms. This is what Archi had to say about this:

Most of these personal names belong to a name-giving tradition different from that of Ebla; Arra-ti/tulu(m) is attested also at Dulu, a neighbouring city-state (Bonechi 1990b: 22–25).28 We must, therefore, deduce that Armi belonged to a marginal, partially Semitized linguistic area different from the ethno-linguistic region dominated by Ebla. Typical are masculine personal names ending in -a-du: A-la/li-wa-du/da, A-li/lu-wa-du, Ba-mi-a-du, La-wadu, Mi-mi-a-du, Mu-lu-wa-du. This reminds one of the suffix -(a)nda, -(a)ndu, very productive in the Anatolian branch of Indo-European (Laroche 1966: 329). Elements such as ali-, alali-, lawadu-, memi-, mula/i- are attested in Anatolian personal names of the Old Assyrian period (Laroche 1966: 26–27, 106, 118, 120).

First_Eblaite_Empire
Ebla’ first kingdom at its height c. 2340 BC. Hipothetical location of Armi depicted. The first Eblaite kingdom extended from Urshu in the north,1 to Damascus area in the south.2 And from Phoenicia and the coastal mountains in the west,3 4 to Tuttul,5 and Haddu in the east.6 The eastern kingdom of Nagar controlled most of the Khabur basin from the river junction with the Euphrates to the northwestern part at Nabada.7 Page 101. From Wikipedia.

This was used by Archi to speculatively locate the state of Armi, in or near Ebla territory, which could correspond with the region of modern north-western Syria:

The onomastic tradition of Armi, so different from that of Ebla and her allies (§ 5), obliges us to locate this city on the edges of the Semitized area and, thus, necessarily north of the line running through Hassuwan – Ursaum – Irritum – Harran. If Armi were to be found at Banat-Bazi, it would have represented an anomaly within an otherwise homogenous linguistic scenario.34

Taken as a whole, the available information suggests that Armi was a regional state, which enjoyed a privileged relationship with Ebla: the exchange of goods between the two cities was comparable only to that between Ebla and Mari. No other state sent so many people to Ebla, especially merchants, lú-kar. It is only a hypothesis that Armi was the go-between for Ebla and for the areas where silver and copper were extracted.

This proposal is similar to the one used to support Indo-Aryan terminology in Mittanni (ca. 16th-14th c. BC), so the scarce material should not pose a problem to those previously arguing about the ‘oldest’ nature of Indo-Aryan.

NOTE. On the other hand, the theory connecting ‘mariannu‘, a term dated to 1761 BC (referenced also in the linguistic supplement), and put in relation with PIIr. *arya, seems too hypothetical for the moment, although there is a clear expansion of Aryan-related terms in the Middle East that could support one or more relevant eastern migration waves of Indo-Aryans from Asia.

Potential routes of Anatolian migration

Once we have accepted that Anatolian is not Late PIE – and that only needed a study of Anatolian archaisms, not the terminology from Armi – , we can move on to explore the potential routes of expansion.

On the Balkan route

A current sketch of the dots connecting Khvalynsk with Anatolia is as follows.

suvorovo-scepters
1—39 — sceptre bearers of the type Giurgiuleşti and Suvorovo; 40—60 — Gumelniţa-Varna-Bolgrad-Aldeni cultural sphere; 61 — Fălciu; 62 — Cainari; 63 — Giurgiuleşti; 64 — Suvorovo; 65 — Casimcea; 66 — Kjulevča; 67 — Reka Devnja; 68 — Drama; 69 — Gonova Mogila; 70 — Reževo.

First, we have the early expansion of Suvorovo chieftains spreading from ca. 4400-4000 BC in the lower Danube region, related to Novodanilovka chiefs of the North Pontic region, and both in turn related to Khvalynsk horse riders (read a a recent detailed post on this question).

Then we have Cernavoda I (ca. 3850-3550 BC), a culture potentially derived from the earlier expansion of Suvorovo chiefs, as shown in cultural similarities with preceding cultures and Yamna, and also in the contacts with the North Pontic steppe cultures (read a a recent detailed post on this question).

We also have proof of genetic inflow from the steppe into populations of cultures near those suggested to be heirs of those dominated by Suvorovo chiefs, from the 5th millennium BC (in Varna I ca. 4630 BC, and Smyadovo ca. 4500 BC, see image below).

If these neighbouring Balkan peoples of ca. 4500 BC are taken as proxies for Proto-Anatolians, then it becomes quite clear why Old Hittite samples dated 3,000 years after this migration event of elite chiefs could show no or almost no ancestry from Europe (for this question, read my revision of Lazaridis’ preprint).

NOTE. A full account of the crisis in the lower Danube, as well as the Suvorovo-Novodanilovka intrusion, is available in Anthony (2007).

mathieson-2018-balkan-expansion
Modified image, including PCA and supervised ADMIXTURE data from Mathieson et al. (2018). Blue arrow represents incoming ancestry from Suvorovo chiefs, red line represents distance from the majority of the neighbouring Balkan population in this period studied to date. Northwestern-Anatolian Neolithic (grey), Yamnaya from Samara (yellow), EHG (pink) and WHG (green).

The southern Balkans and Anatolia

The later connection of Cernavoda II-III and related cultures (and potentially Ezero) with Troy, on the other hand, is still blurry. But, even if a massive migration of Common Anatolian is found to happen from the Balkans into Anatolia in the late 4th / beginning of the 3rd millennium, the people responsible for this expansion could show a minimal trace of European ancestry.

A new paper has appeared recently (in Russian), Dubene and Troy: Gold and Prosperity in the Third Millennium Cal. BCE in Eurasia. Stratum Plus, 2 (2018), by L. Nikolova, showing commercial contacts between Troy and cultures from Bulgaria:

Earlier third millennium cal BCE is the period of development of interconnected Early Bronze Age societies in Eurasia, which economic and social structures expressed variants of pre-state political structures, named in the specialized literature tribes and chiefdoms. In this work new arguments will be added to the chiefdom model of third millennium cal BC societies of Yunatsite culture in the Central Balkans from the perspectives of the interrelations between Dubene (south central Bulgaria) and Troy (northwest Turkey) wealth expression.

Possible explanations of the similarity in the wealth expression between Troy and Yunatsite chiefdoms is the direct interaction between the political elite. However, the golden and silver objects in the third millennium cal BCE in the Eastern Mediterranean are most of all an expression of economic wealth. This is the biggest difference between the early state and chiefdoms in the third millennium cal BCE in Eurasia and Africa. The literacy and the wealth expression in the early states was politically centralized, while the absence of literacy and wider distribution of the wealth expression in the chiefdoms of the eastern Mediterranean are indicators, that wider distribution of wealth and the existed stable subsistence layers prevented the formation of states and the need to regulate the political systems through literacy.

The only way to link Common Anatolians to their Proto-Anatolian (linguistic) ancestors would therefore be to study preceding cultures and their expansions, until a proper connecting route is found, as I said recently.

These late commercial contacts in the south-eastern Balkans (Nikolova also offers a simplified presentation of data, in English) are yet another proof of how Common Anatolian languages may have further expanded into Anatolia.

NOTE. One should also take into account the distribution of modern R1b-M269* and L23* subclades (i.e. those not belonging to the most common subclades expanding with Yamna), which seem to peak around the Balkans. While those may just belong to founder effects of populations preceding Suvorovo or related to Yamna migrants, the Balkans is a region known to have retained Y-DNA haplogroup diversity, in contrast with other European regions.

On a purely linguistic aspect, there are strong Hattic and Hurrian influences on Anatolian languages, representing a unique layer that clearly differentiates them from LPIE languages, pointing also to different substrates behind each attested Common Anatolian branch or individual language:

  • Phonetic changes, like the appearance of /f/ and /v/.
  • Split ergativity: Hurrian is ergative, Hattic probably too.
  • Increasing use of enclitic pronoun and particle chains after first stressed word: in Hattic after verb, in Hurrian after nominal forms.
  • Almost obligatory use of clause initial and enclitic connectors: e.g. semantic and syntactic identity of Hattic pala/bala and Hittite nu.

NOTE. For a superficial discussion of this, see e.g. An Indo-European Linguistic Area and its Characteristics: Ancient Anatolia. Areal Diffusion as a Challenge to the Comparative Method?, by Calvert Watkins. You can also search for any of the mentioned shared isoglosses between Middle Eastern languages and Anatolian if you want more details.

On the Caucasus route

It seems that the Danish group is now taking a stance in favour of a Maykop route (from the linguistic supplement):

The period of Proto-Anatolian linguistic unity can now be placed in the 4th millennium BCE and may have been contemporaneous with e.g. the Maykop culture (3700–3000 BCE), which influenced the formation and apparent westward migration of the Yamnaya and maintained commercial and cultural contact with the Anatolian highlands (Kristiansen et al. 2018).

In fact, they have data to support this:

The EHG ancestry detected in individuals associated with both Yamnaya (3000–2400 BCE) and the Maykop culture (3700–3000 BCE) (in prep.) is absent from our Anatolian specimens, suggesting that neither archaeological horizon constitutes a suitable candidate for a “homeland” or “stepping stone” for the origin or spread of Anatolian Indo- European speakers to Anatolia. However, with the archaeological and genetic data presented here, we cannot reject a continuous small-scale influx of mixed groups from the direction of the Caucasus during the Chalcolithic period of the 4th millennium BCE.

While it is difficult to speak about the consequences of this find without having access to this paper in preparation or its samples, we already knew that Maykop had obvious cultural contacts with the steppe.

It will not be surprising to find not only EHG, but also R1b-L23 subclades there. In my opinion, though, the most likely source of EHG ancestry in Maykop (given the different culture shown in other steppe groups) is exogamy.

The question will still remain: was this a Proto-Anatolian-speaking group?

eneolithic_steppe
Diachronic map of Eneolithic migrations ca. 4000-3100 BC

My opinion in this regard – again, without access to the study – is that you would still need to propose:

  • A break-up of Anatolian ca. 4500 BC represented by some early group migrating into the Northern Caucasus area.
  • For this group – who were closely related linguistically and culturally to early Khvalynsk – to remain isolated in or around the Northern Caucasus, i.e. somehow ‘hidden’ from the evolving LPIE speakers in late Khvalynsk/early Yamna peoples.
  • Then, they would need to have migrated from Maykop to Anatolian territory only after ca. 3700 BC – while having close commercial contacts with Khvalynsk and the North Pontic cultures in the period 3700-3000 BC -, in some migration wave that has not showed up in the archaeological records to date.
  • Then appear as Old Hittites without showing EHG ancestry (even though they show it in the period 3700-3000 BC), near the region of the Armi state, where Anatolian was supposedly spoken already in the mid-3rd millennium.

Not a very convincing picture, right now, but indeed possible.

Also, we have R1b-Z2103 lineages and clear steppe ancestry in the region probably ca. 2500 BC with Hajji Firuz, which is most likely the product of the late Khvalynsk migration waves that we are seeing in the recent papers.

These migrations are then related to early LPIE-speaking migrants spreading after ca. 3300 BC – that also caused the formation of early Yamna and the expansion of Tocharian-related migrants – , which leaves almost no space for an Anatolian expansion, unless one supports that the former drove the latter.

NOTE. In any case, if the Caucasus route turned out to be the actual Anatolian route, I guess this would be a way as good as any other to finally kill their Indo-European – Corded Ware theory, for obvious reasons.

On the North Iranian homeland

A few thoughts for those equating CHG ancestry in IE speakers (and especially now in Old Hittites) with an origin in North Iran, due to a recent comment by David Reich:

In the paper it is clearly stated that there is no Neolithic Iranian ancestry in the Old Hittite samples.

Ancestry is not people, and it is certainly not language. The addition of CHG ancestry to the Eneolithic steppe need not mean a population or linguistic replacement. Although it could have been. But this has to be demonstrated with solid anthropological models.

NOTE. On the other hand, if you find people who considered (at least until de Barros Damgaard et al. 2018) steppe (ancestry/PCA) = Indo-European, then you should probably confront them about why CHG in Hittites and the arrival of CHG in steppe groups is now not to be considered the same, i.e why CHG / Iran_N ≠ PIE.

Since there has been no serious North Iranian homeland proposal made for a while, it is difficult to delineate a modern sketch, and I won’t spend the time with that unless there is some real anthropological model and genetic proof of it. I guess the Armenian homeland hypothesis proposed by Gamkrelidze and Ivanov (1995) would do, but since it relies on outdated data (some of which appears also in Gimbutas’ writings), it would need a full revision.

NOTE. Their theory of glottalic consonants (or ejectives) relied on the ‘archaism’ of Hittite, Germanic, and Armenian. As you can see (unless you live in the mid-20th century) this is not very reasonable, since Hittite is attested quite late and after heavy admixture with Middle Eastern peoples, and Germanic and Armenian are some of the latest attested (and more admixed, phonetically changed) languages.

This would be a proper answer, indeed, for those who would accept this homeland due to the reconstruction of ‘ejectives’ for these languages. Evidently, there is no need to posit a homeland near Armenia to propose a glottalic theory. Kortlandt is a proponent of a late and small expansion of Late PIE from the steppe, and still proposes a reconstruction of ejectives for PIE. But, this was the main reason of Gamkrelidze and Ivanov to propose that homeland, and in that sense it is obviously flawed.

Those claiming a relationship of the North Iranian homeland with such EHG ancestry in Maykop, or with the hypothetic Proto-Euphratic or Gutian, are obviously not understanding the implications of finding steppe ancestry coupled with (likely) early Late PIE migrants in the region in the mid-4th millennium.

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Consequences of O&M 2018 (II): The unsolved nature of Suvorovo-Novodanilovka chiefs, and the route of Proto-Anatolian expansion

neolithic_steppe-suvorovo

This is part of a series of posts analyzing the findings of the recent Nature papers Olalde et al.(2018) and Mathieson et al.(2018) (abbreviated O&M 2018).

I already expressed my predictions for 2018. One of the most interesting questions among them is the identification of the early Anatolian offshoot, and this is – I believe – where Genomics has the most to say in Indo-European migrations.

Linguistics and Archaeology had already a mainstream account from Late PIE/Yamna onwards, and it has been proven right in Genomic investigation. There is, however, no consensus on Indo-Hittite.

Suvorovo-Novodanilovka

Apart from the Anatolian homeland hypothesis and its westward migration (as referenced e.g. by Lazaridis et al. 2017), the other possibility including the most likely steppe homeland is that Proto-Anatolian spread through the Balkans, and must have separated from Khvalynsk and travelled first westward through the North Pontic region, and then southward to Ezero.

EDIT (10 MAR 2018): The Anatolian westward route within the steppe homeland model refers to the possibility that Proto-Anatolian spread south through the Caucasus, and then westward through Anatolia, as suggested e.g. originally by Marija Gimbutas for Maykop, as a link in the Caucasus.

We all know that this Khvalynsk -> Novodanilovka-Suvorovo -> Cernavoda -> Ezero -> Troy migration model proposed by Anthony shows no conspicuous chain in Archaeology, but obvious contacts (including Genomics) are seen among some of these neighbouring cultures in different times.

We know that remains of Suvorovo-Novodanilovka culture of chiefs emerged around 4400-4200 BC among ordinary local Sredni Stog settlements:

  • the Novodanilovka rich burials in the steppes, near the Dnieper,
  • and the Suvorovo group in the Danube delta, roughly coinciding with the massive abandonment of old tell settlements in the area.

One of the strongest cultural connections between Khvalynsk and Suvorovo Novodanilovka chiefs is the similar polished stone mace-heads shaped like horse heads found in both cultures, a typical steppe prestige object going back to the east Pontic-Caspian steppe beginning ca. 5000-4800 BC.

Its finding in the Danube valley may have signalled the expansion of horse riding, which is compatible with the finding of ancient domesticated horses in the region. Horses were not important in Old European cultures, and it seems that they weren’t in Sredni Stog or Kvitjana either.

sredni-stog-suvorovo-novodanilovka-cernavoda
Steppe and Danubian sites at the time: of the Suvorovo-Novodanilovka intrusion, about 4200-3900 BC. David W. Anthony (2007).

NOTE. Telegin, the main source of knowledge in Ukraine prehistoric cultures for Anthony, was eventually convinced that Surovovo-Novodanilovka was a separate culture. However, for Anthony (using Telegin’s first impressions), it may have been a wealthy elite among Sredni Stog peoples. Anthony considers Sredni Stog to have been also influenced by Khvalynsk, and thus potentially related to the Suvorovo-Novodanilovka chiefs.

Nevertheless, he obviously cannot link North Pontic Eneolithic cultures to Khvalynsk nor to horse riding – whilst he clearly assumes horse riding for Novodanilovka-Suvorovo chiefs – , and he does not link North Pontic cultures to later expansions of Late Proto-Indo-Europeans from late Khvalynsk and Yamna, either.

The question here for Anthony (as with further Proto-Anatolian expansions described in his 2007 book), in my opinion, was to offer a plausible string of connections between Khvalynsk and Anatolia, and the simplest connection one can make among steppe cultures is a general, broad community between North Pontic and North Caspian cultures. That way, the knot tying Khvalynsk to the Danube seems stronger, whatever the origin of Suvorovo-Novodanilovka chiefs.

If, however, a direct genetic connection is made between Suvorovo-Novodanilovka chiefs and Khvalynsk – as in its association with R1b-M269 and R1b-L23 lineages – , there will be little need to include Sredni Stog or any other intermediate culture in the equation.

We have already seen a movement of steppe ancestry into mainland Greece, and I would not be surprised if a parallel movement could be seen from Ezero to Troy (or a neighbouring North-West Anatolian region), so that the final migration of Common Anatolian had in fact been triggered by the massive steppe migrations during the Chalcolithic.

NOTE. Whereas we are certain to find R1b-L23 subclades in the direct Balkan migrations from Yamna, the link of steppe->Anatolia migrations may be a little trickier: even if we find out that the Suvorovo-Novodanilovka expansion was associated with an expansion and reduction of haplogroup variability (to haplogroups R1b-M269 and R1b-L23), we don’t know yet if the ca. 1,500 years passed (and the different cultural and population changes occurred) between Proto-Anatolian and Common Anatolian migrations may have impacted the main haplogroup composition of both communities.

O&M 2018

A probably unsurprising – because of its previously known admixture and PCA – , but nevertheless disappointing finding came from the Y-SNP call of the haplogroup R1 found in Varna (R1b-V88, given first by Genetiker), leaving us with no new haplogroup data standing out for this period.

This sample’s lack of obvious genetic links with the steppe and early date didn’t deter me from believing it could show subclade M269, and thus a sign of incoming Suvorovo chiefs in the region. After all, R1b-P297 subclades seemed to have almost disappeared from the Balkans by that time, and we know that assessments based only on ancestral components and PCA clusters are not infallible – we are seeing that in many, many samples already.

suvorovo-scepters
1—39 — sceptre bearers of the type Giurgiuleşti and Suvorovo; 40—60 — Gumelniţa-Varna-Bolgrad-Aldeni cultural sphere; 61 — Fălciu; 62 — Cainari; 63 — Giurgiuleşti; 64 — Suvorovo; 65 — Casimcea; 66 — Kjulevča; 67 — Reka Devnja; 68 — Drama; 69 — Gonova Mogila; 70 — Reževo. Țerna S., Govedarica B. (2016)

NOTE. In fact, the first time I checked Mathieson et al. (2018) supplementary tables I thought that the ‘Ukraine_Eneolithic’ sample of R1b-L23 subclade was ‘it’: the first clear proof in ancient samples of incoming Suvorovo chiefs from Khvalynsk I was looking for…Until I realized its date, and that it was more likely a Late Yamna (or Catacomb) sample.

Steppe ancestry is found in the Varna and Smyadovo outliers, though, and these samples cluster closely to Ukraine Eneolithic samples (which are among Khvalynsk, Ukraine Neolithic, and Anatolia Neolithic clusters), so some population movement must have happened around or before that time in the region, and it is obvious that it happened from east to west.

It remains to be seen, therefore:

a) If the incoming Suvorovo-Novodanilovka chiefs (most likely originally from Khvalynsk) dominating over North Pontic and Danube regions show – as I bet – R1b-M269, and possibly also early R1b-L23* subclades,

b) Or else they still show mixed lineages, reflecting an older admixed population of the Pontic-Caspian steppe – as the early Khvalynsk and Ukraine Eneolithic samples we have now.

NOTE. Even though my preferred model of migration is through the Balkans – due to the many east-west migrations seen from the steppe into Europe – , there is no general consensus here because of the lack of solid anthropological models, and there are cultural links found also between the steppe and Anatolia through the Caucasus, so the question remains open.

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