First Iberian R1b-DF27 sample, probably from incoming East Bell Beakers


I had some more time to read the paper by Valdiosera et al. (2018) and its supplementary material.

One of the main issues since the publication of Olalde et al. (2018) (and its hundreds of Bell Beaker samples) was the lack of a clear Y-DNA R1b-DF27 subclades among East Bell Beaker migrants, which left us wondering when the subclade entered the Iberian Peninsula, since it could have (theoretically) happened from the Chalcolithic to the Iron Age.

My prediction was that this lineage found today widespread among the Iberian population crossed the Pyrenees quite early, during the Chalcolithic, with migrating East Bell Beakers expanding North-West Indo-European dialects, and that it spread slowly afterwards.

The first ancient sample clearly identified as of R1b-DF27 subclade is found in this paper, at the Late Bronze Age site Cueva de los Lagos. Although it is unidentified and has no radiocarbon date, the site as a whole is associated with the Cogotas culture and its Bouquique ceramic decoration.

Y-DNA and mtDNA haplogroups, from the paper. Sequencing statistics and contamination rates for newly generated sequence data.

It was found in the northern part of the Cogotas culture territory (which lies mainly between Castille and Aragon, in North-Central Spain), shows evident steppe admixture, and it has become obvious with the latest papers (including this one) that R1b-M269 lineages intruded south of the Pyrenees associated with East Bell Beaker migrations.

The Proto-Cogotas culture is associated with a Bell Beaker substrate influenced by either El Argar or Atlantic Bronze, and the specific type of ceramics found at this Cogotas culture site are probably from the mid-2nd millennium, which is too early for the Celtic expansion.

Supervised ADMIXTURE results.

Nevertheless, due to the quite likely late date of the sample (in the centuries around 1500 BC), there is still a possibility that incoming R1b-DF27 lineages were not among the early R1b-M269 lineages found in the Iberian Chalcolithic, and were associated with later migrations from Central Europe, potentially linked to the expansion of the Urnfield culture, and thus nearer to an Italo-Celtic community.

Diachronic map of migrations in Europe ca. 1250-750 BC.

In any of these scenarios, a Pre-Celtic expansion of North-West Indo-European in Iberia (possibly associated with Lusitanian) is still the best explanation for the origin and expansion of (at least some) modern Iberian R1b-DF27 lineages, including those found among the Basque-speaking population.

This implies that the ‘indigenous’ Neolithic lineages of Iberia (like I2 and G2a2) were replaced with subsequent internal gene flows and founder effects, such as those that evidently happened (probably quite recently) among Basques, even though indigenous languages show an obvious continuity.

I would say this is the last nail in the coffin for autochthonous Y-DNA continuity theories for Spain and France (i.e. for the traditional Vasconic-Uralic hypothesis), but we know that data is never enough for any die hard continuist…so let’s just say another nail in the coffin for endless autochthonous continuity theories.

EDIT (18/3/2018): Genetiker has published Y-SNP calls for both R1b samples, showing this one is R1b1a1a2a1a2a-BY15964 (see modern members of this subclade in ytree), and that the other one is R1b1a1a2a~L23.


Iberian prehistoric migrations in Genomics from Neolithic, Chalcolithic, and Bronze Age


New open access paper Four millennia of Iberian biomolecular prehistory illustrate the impact of prehistoric migrations at the far end of Eurasia, by Valdiosera, Günther, Vera-Rodríguez, et al. PNAS (2018) published ahead of print.

Abstract (emphasis mine)

Population genomic studies of ancient human remains have shown how modern-day European population structure has been shaped by a number of prehistoric migrations. The Neolithization of Europe has been associated with large-scale migrations from Anatolia, which was followed by migrations of herders from the Pontic steppe at the onset of the Bronze Age. Southwestern Europe was one of the last parts of the continent reached by these migrations, and modern-day populations from this region show intriguing similarities to the initial Neolithic migrants. Partly due to climatic conditions that are unfavorable for DNA preservation, regional studies on the Mediterranean remain challenging. Here, we present genome-wide sequence data from 13 individuals combined with stable isotope analysis from the north and south of Iberia covering a four-millennial temporal transect (7,500–3,500 BP). Early Iberian farmers and Early Central European farmers exhibit significant genetic differences, suggesting two independent fronts of the Neolithic expansion. The first Neolithic migrants that arrived in Iberia had low levels of genetic diversity, potentially reflecting a small number of individuals; this diversity gradually increased over time from mixing with local hunter-gatherers and potential population expansion. The impact of post-Neolithic migrations on Iberia was much smaller than for the rest of the continent, showing little external influence from the Neolithic to the Bronze Age. Paleodietary reconstruction shows that these populations have a remarkable degree of dietary homogeneity across space and time, suggesting a strong reliance on terrestrial food resources despite changing culture and genetic make-up.

(A) f4 statistics testing affinities of prehistoric European farmers to either early Neolithic Iberians or central Europeans, restricting these reference populations to SNP-captured individuals to avoid technical artifacts driving the affinities. The boxplots in A show the distributions of all individual f4 statistics belonging to the respective groups. The signal is not sensitive to the choice of reference populations and is not driven by hunter-gatherer–related admixture (Datasets S4 and S5). (B) Estimates of ancestry proportions in different prehistoric Europeans as well as modern southwestern Europeans. Individuals from regions of Iberia were grouped together for the analysis in A and B to increase sample sizes per group and reduce noise


We present a comprehensive biomolecular dataset spanning four millennia of prehistory across the whole Iberian Peninsula. Our results highlight the power of archaeogenomic studies focusing on specific regions and covering a temporal transect. The 4,000 y of prehistory in Iberia were shaped by major chronological changes but with little geographic substructure within the Peninsula. The subtle but clear genetic differences between early Neolithic Iberian farmers and early Neolithic central European farmers point toward two independent migrations, potentially originating from two slightly different source populations. These populations followed different routes, one along the Mediterranean coast, giving rise to early Neolithic Iberian farmers, and one via mainland Europe forming early Neolithic central European farmers. This directly links all Neolithic Iberians with the first migrants that arrived with the initial Mediterranean Neolithic wave of expansion. These Iberians mixed with local hunter-gatherers (but maintained farming/pastoral subsistence strategies, i.e., diet), leading to a recovery from the loss of genetic diversity emerging from the initial migration founder bottleneck. Only after the spread of Bell Beaker pottery did steppe-related ancestry arrive in Iberia, where it had smaller contributions to the population compared with the impact that it had in central Europe. This implies that the two prehistoric migrations causing major population turnovers in central Europe had differential effects at the southwestern edge of their distribution: The Neolithic migrations caused substantial changes in the Iberian gene pool (the introduction of agriculture by farmers) (6, 9, 11, 13, 24), whereas the impact of Bronze Age migrations (Yamnaya) was significantly smaller in Iberia than in north-central Europe (24). The post-Neolithic prehistory of Iberia is generally characterized by interactions between residents rather than by migrations from other parts of Europe, resulting in relative genetic continuity, while most other regions were subject to major genetic turnovers after the Neolithic (4, 6, 7, 9, 25, 48). Although Iberian populations represent the furthest wave of Neolithic expansion in the westernmost Mediterranean, the subsequent populations maintain a surprisingly high genetic legacy of the original pioneer farming migrants from the east compared with their central European counterparts. This counterintuitive result emphasizes the importance of in-depth diachronic studies in all parts of the continent.


The origin and expansion of Pama–Nyungan languages across Australia

Yet another questionable paper by Nature, The origin and expansion of Pama–Nyungan languages across Australia, by Bouckaert, Bowern & Atkinson, Nat Ecol Evol (2018).


It remains a mystery how Pama–Nyungan, the world’s largest hunter-gatherer language family, came to dominate the Australian continent. Some argue that social or technological advantages allowed rapid language replacement from the Gulf Plains region during the mid-Holocene. Others have proposed expansions from refugia linked to climatic changes after the last ice age or, more controversially, during the initial colonization of Australia. Here, we combine basic vocabulary data from 306 Pama–Nyungan languages with Bayesian phylogeographic methods to explicitly model the expansion of the family across Australia and test between these origin scenarios. We find strong and robust support for a Pama–Nyungan origin in the Gulf Plains region during the mid-Holocene, implying rapid replacement of non-Pama–Nyungan languages. Concomitant changes in the archaeological record, together with a lack of strong genetic evidence for Holocene population expansion, suggests that Pama–Nyungan languages were carried as part of an expanding package of cultural innovations that probably facilitated the absorption and assimilation of existing hunter-gatherer groups.

“Diversification of the Pama–Nyungan language family. Maximum clade credibility tree showing the inferred timing and emergence of the major branches and their subsequent diversification.”

Even with my absolute lack of knowledge on Australian languages, I am not conviced. Not at all.

I have already expressed more than once my opinion on Glottochronology – and the improved method of this paper seems like the final twist of the screw for its strongest proponents.

Interestingly, this paper includes the same journal, author, and (mostly) method of the famous Language-tree divergence times support the Anatolian theory of Indo-European origin (2003).

And we have also seen how most suggested prehistorical cultural diffusion events were actually migrations, so it seems rather odd to dare publish this right now.

At a time of groundbreaking genomic papers being published on South-East Asian migrations, and probably expecting more on the region – including Australia – , this paper seems to me quite unnecessary.

It will especially not help Nature make forget its latest fiasco on Indo-European migrations.

See also:

On the potential origin of Caucasus hunter-gatherer ancestry in Eneolithic steppe cultures

An interesting open genomic question is the origin and spread of Caucasus hunter-gatherer (CHG) ancestry in steppe populations during the Eneolithic.

My broad theory regarding the appearance of this ancestral component is based on:

Two recently published papers ivestigating the Don Region may shed some light on this issue:

Plant food subsistence in the human diet of the Bronze Age Caspian and Low Don steppe pastoralists: archaeobotanical, isotope and 14C data, by Shishlina, Bobrov, Simakova, et al. Veget Hist Archaeobot (2018).

EDIT (16/3/2018): You can now read or download the paper at


The paper presents the result of analysis of charred food on the interior part of the vessels from the graves of the East Manych and West Manych Catacomb archaeological cultures (2500–2350 cal bc). The phytolith and pollen analyses identified pollen of wild steppe plants and phytoliths of domesticated gramineous plants determined as barley phytoliths. Direct 14С dating of one of the samples demonstrates that barley spikelets and stems were used in funeral rites by local steppe communities. However, there are no data suggesting that steppe inhabitants of the Lower Don Region were engaged in agriculture in the mid-3000 bc. Supposedly, barley could have reached the steppes through seasonal migrations of mobile pastoralists to the south, use of North Caucasus grasslands in the economic system of seasonal moves and exchange with local people. Nevertheless, presence of carbonized barley seeds in the occupation layers at North Caucasus settlements of 4000–3000 bc requires confirmation by direct 14С dating of such samples.

Location of sites. 1: Ulan IV; 2: Peschany IV and V; 3: Shakhaevskaya 1; 4: Zunda-Tolga 2; 5: Lesnoye; 6: Chidgom; 7: Meshoko; 8: Chishkho; 9: Svobodnoye

Dynamics of Chemical and Microbiological Soil Properties in the Desert–Steppe Zone of the Southeast Russian Plain during the Second Part of the Holocene (4000 BC–XIII century AC), Kashirskaya, Khomutova, Kuznetsova, et al. Arid Ecosyst (2018) 8(1):38-46.


The results of studies of the chemical and microbiological properties of the soils buried under the barrows of the Eneolithic, Bronze, and Middle Ages periods of the southeast of the Russian Plain are presented. It was shown that the climate of the region in the Eneolithic period (4200–4100 BC) and in the Middle Ages (700 years ago) was more humid in comparison to the present time. The third millennium BC was characterized by a gradual increase of the climate aridity. Its peak was at the end of the III millennium BC. The number and biomass of microbial cells was maximal in soils buried in periods of high atmospheric humidity (4200–4100 and 3000–2800 BC) and sharply decreased during the aridization period in the second half of the III millennium BC. In general, the variability of indicators of microbocenosis conditions of desert–steppe buried soils of all ages from the burial mounds correlated with the centuries-old dynamics of the climate.

Number of microbial cells in buried soils of different ages and modern background soil.

It is well known that access to more food – as in favorable crops and cattle feeding – may cause demographic explosions, and the second article – together with recent genomic data – may be yet another proof of that.

Until now, pastoralism seemed to be the main subsistence economy for most steppe groups. It seems that earlier Eneolithic contacts of certain steppe groups with settlements of the Northern Caucasus might have been not just to obtain prestige goods though, but – if proper radiocarbon dating confirms it – also implied essential goods, and maybe more stable seasonal exchange systems.

Such stable economic exchanges might have therefore included bidirectional exogamy practices, justifying the sizeable genomic contribution from the Caucasus.

At this point this is just another good theory to take into account.


Genomic analysis of Germanic tribes from Bavaria show North-Central European ancestry


New open access paper Population genomic analysis of elongated skulls reveals extensive female-biased immigration in Early Medieval Bavaria, by Veeramah, Rott, Groß, et al. PNAS (2018), published ahead of print.

First, a bit of context on the Bavarii:

Europe experienced a profound cultural transformation between Late Antiquity and the Middle Ages that laid the foundations of the modern political, social, and religious landscape. During this period, colloquially known as the “Migration Period,” the Roman Empire gradually dissolved, with 5th and 6th century historiographers and contemporary witnesses describing the formation and migration of numerous Germanic peoples, such as the Goths, Alamanni, Gepids, and Longobards. However, the genetic and social composition of groups involved and the exact nature of these “migrations” are unclear and have been a subject of substantial historical and archaeological debate

In the mid 6th century AD, the historiographer Jordanes and the poet and hagiographer Venantius Fortunatus provide the first mention of a group known as the Baiuvarii that resided in modern day Bavaria. It is likely that this group had already started to form in the 5th century AD, and that it emanated from a combination of the romanized local population of the border province of the former Roman Empire and immigrants from north of the Danube (2). While the Baiuvarii are less well known than some other contemporary groups, an interesting archaeological feature in Bavaria from this period is the presence of skeletons with artificially deformed or elongated skulls.

Procrustes-transformed PCA of ancient samples using pseudohaploid calls based on off-target reads using an imputed POPRES modern reference dataset. Blue, green, and red male or female symbols are ancient Bavarian individuals with normal, intermediate, and elongated skulls, respectively. Orange circles are Anglo-Saxon era individuals. Large circles are medians for regions, dots are individuals. CE, central Europe; EE, eastern Europe; NE, northern Europe; NEE, northeastern Europe; NEW, northwestern Europe; SE, southern Europe; SEE, southeast Europe; WE, western Europe. Percentage of variation explained by PCs 1 and 2 for modern populations only is 0.25% and 0.15%.

Abstract (emphasis mine):

Modern European genetic structure demonstrates strong correlations with geography, while genetic analysis of prehistoric humans has indicated at least two major waves of immigration from outside the continent during periods of cultural change. However, population-level genome data that could shed light on the demographic processes occurring during the intervening periods have been absent. Therefore, we generated genomic data from 41 individuals dating mostly to the late 5th/early 6th century AD from present-day Bavaria in southern Germany, including 11 whole genomes (mean depth 5.56×). In addition we developed a capture array to sequence neutral regions spanning a total of 5 Mb and 486 functional polymorphic sites to high depth (mean 72×) in all individuals. Our data indicate that while men generally had ancestry that closely resembles modern northern and central Europeans, women exhibit a very high genetic heterogeneity; this includes signals of genetic ancestry ranging from western Europe to East Asia. Particularly striking are women with artificial skull deformations; the analysis of their collective genetic ancestry suggests an origin in southeastern Europe. In addition, functional variants indicate that they also differed in visible characteristics. This example of female-biased migration indicates that complex demographic processes during the Early Medieval period may have contributed in an unexpected way to shape the modern European genetic landscape. Examination of the panel of functional loci also revealed that many alleles associated with recent positive selection were already at modern-like frequencies in European populations ∼1,500 years ago.

Supervised model-based clustering ADMIXTURE analysis for ancient samples based on phased haplotypes for individual 1,000 bp loci from the 5-Mb neutralome. Analysis is based on the best of 100 runs for K = 8, but NC_EUR is the ancestry summed across 1000 Genomes CEU, 1000 Genomes GBR, and GoNL populations (i.e., it represents a northern/central European ancestry). Blue, green, and red male or female symbols are ancient Bavarian individuals with normal, intermediate, and elongated skulls, respectively.

There is no Y-DNA data to keep confirming the North-Central origin of certain modern European subclades in Central and South-Central Europe.

The potential Ostrogothic sample from Crimea was probably Hunnic, as the paper itself suggests, and both Ostrogoths and Gepids are known to have been allies of the Huns for a long time. It is also a well-known fact that East Germanic tribes migrated south- and eastward through eastern Europe, and then from the steppe westward.

Obviously, the PCA of a late Gepid sample – after a certain number of generations and admixture events with ‘local’ populations during the migrations – , and of a Crimean sample without a clear cultural identification, are of limited value today, until more samples are available.

Hence sadly no valid data yet to add to the debate of East Germanic nature, which mainly concerns its traditionally described origin in Scandinavia – i.e. close to North Germanic dialects – against a different origin (and dialectal branch) within Proto-Germanic territory.

NOTE. Just to be clear for future papers on Germanic tribes, I would expect East Germanic males to show either:
a) mainly R1b-U106, I1, and R1a-Z645 subclades, and to cluster closely to samples of Scandinavia during Antiquity, which would support a Scandinavian origin – a predominance of typically Scandinavian R1a-Z284 subclades would be more indicative of this origin, of course;
b) or mainly R1b-U106, R1b-P312, and I1 subclades and a PCA cluster close to West Germanic tribes, which would challenge its traditional dialectal identification.

I agree with the authors in that a few samples are able to describe certain migratory events, though, such as the emphasized female-biased long-distance migration in Bavaria, as well as the diverse ancestry of women versus men.


Patterns of genetic differentiation and the footprints of historical migrations in the Iberian Peninsula

Open access preprint (which I announced already) at bioRxiv Patterns of genetic differentiation and the footprints of historical migrations in the Iberian Peninsula, by Bycroft et al. (2018).

Abstract (emphasis mine):

Genetic differences within or between human populations (population structure) has been studied using a variety of approaches over many years. Recently there has been an increasing focus on studying genetic differentiation at fine geographic scales, such as within countries. Identifying such structure allows the study of recent population history, and identifies the potential for confounding in association studies, particularly when testing rare, often recently arisen variants. The Iberian Peninsula is linguistically diverse, has a complex demographic history, and is unique among European regions in having a centuries-long period of Muslim rule. Previous genetic studies of Spain have examined either a small fraction of the genome or only a few Spanish regions. Thus, the overall pattern of fine-scale population structure within Spain remains uncharacterised. Here we analyse genome-wide genotyping array data for 1,413 Spanish individuals sampled from all regions of Spain. We identify extensive fine-scale structure, down to unprecedented scales, smaller than 10 Km in some places. We observe a major axis of genetic differentiation that runs from east to west of the peninsula. In contrast, we observe remarkable genetic similarity in the north-south direction, and evidence of historical north-south population movement. Finally, without making particular prior assumptions about source populations, we show that modern Spanish people have regionally varying fractions of ancestry from a group most similar to modern north Moroccans. The north African ancestry results from an admixture event, which we date to 860 – 1120 CE, corresponding to the early half of Muslim rule. Our results indicate that it is possible to discern clear genetic impacts of the Muslim conquest and population movements associated with the subsequent Reconquista.

“(a) Binary tree showing the inferred hierarchical relationships between clusters. The colours and points correspond to each cluster as shown on the map, and the length of the coloured rectangles is proportional to the number of individuals assigned to that cluster. We combined some small clusters (Methods) and the thick black branches indicate the clades of the tree that we visualise in the map. We have labeled clusters according to the approximate location of most of their members, but geographic data was not used in the inference. (b) Each individual is represented by a point placed at (or close to) the centroid of their grandparents’ birthplaces. On this map we only show the individuals for whom all four grandparents were born within 80km of their average birthplace, although the data for all individuals were used in the fineSTRUCTURE inference. The background is coloured according to the spatial densities of each cluster at the level of the tree where there are 14 clusters (see Methods). The colour and symbol of each point corresponds to the cluster the individual was assigned to at a lower level of the tree, as shown in (a). The labels and boundaries of Spain’s Autonomous Communities are also shown.”

Some interesting excerpts:

Our results further imply that north west African-like DNA predominated in the migration. Moreover, admixture mainly, and perhaps almost exclusively, occurred within the earlier half of the period of Muslim rule. Within Spain, north African ancestry occurs in all groups, although levels are low in the Basque region and in a region corresponding closely to the 14th-century ‘Crown of Aragon’. Therefore, although genetically distinct this implies that the Basques have not been completely isolated from the rest of Spain over the past 1300 years.

NOTE. I must add here that the Expulsion of Moriscos is known to have been quite successful in the old Crown of Aragon – deeply affecting its economy – , in contrast with other territories of the Crown of Castille, where they either formed less sizeable communities, or were dispersed and eventually Christened and integrated with local communities. For example, thousands of Moriscos from Granada were dispersed following the War of Alpujarras (1567–1571) into different regions of the Crown of Castille, and many could not be later expelled due to the locals’ resistance to follow the expulsion edict.

Perhaps surprisingly, north African ancestry does not reflect proximity to north Africa, or even regions under more extended Muslim control. The highest amounts of north African ancestry found within Iberia are in the west (11%) including in Galicia, despite the fact that the region of Galicia as it is defined today (north of the Miño river), was never under Muslim rule and Berber settlements north of the Douro river were abandoned by. This observation is consistent with previous work using Y-chromosome data. We speculate that the pattern we see is driven by later internal migratory flows, such as between Portugal and Galicia, and this would also explain why Galicia and Portugal show indistinguishable ancestry sharing with non-Spanish groups more generally. Alternatively, it might be that these patterns reflect regional differences in patterns of settlement and integration with local peoples of north African immigrants themselves, or varying extents of the large-scale expulsion of Muslim people, which occurred post-Reconquista and especially in towns and cities.

We estimated ancestry profiles for each point on a fine spatial grid across Spain (Methods). Gray crosses show
the locations of sampled individuals used in the estimation. Map shows the fraction contributed from the donor group ‘NorthMorocco’.

Overall, the pattern of genetic differentiation we observe in Spain reflects the linguistic and geopolitical boundaries present around the end of the time of Muslim rule in Spain, suggesting this period has had a significant and long-term impact on the genetic structure observed in modern Spain, over 500 years later. In the case of the UK, similar geopolitical correspondence was seen, but to a different period in the past (around 600 CE). Noticeably, in these two cases, country-specific historical events rather than geographic barriers seem to drive overall patterns of population structure. The observation that fine-scale structure evolves at different rates in different places could be explained if observed patterns tend to reflect those at the ends of periods of significant past upheaval, such as the end of Muslim rule in Spain, and the end of the Anglo-Saxon and Danish Viking invasions in the UK.

Certain people want to believe (well into the 21st century) into ideal ancestral populations and ancient ethnolinguistic identifications linked to one’s own – or the own country’s dominant – ancestral components and Y-DNA haplogroup.

We are nevertheless seeing how mainly the most recent relevant geopolitical events and late internal migratory flows have shaped the genetic structure (including Y-DNA haplogroup composition) of modern regions and countries regardless of its population’s actual language or ethnic identification, whether (pre)historical or modern.

Another surprise for many, I guess.


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


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.


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.

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.

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.


North Pontic steppe Eneolithic cultures, and an alternative Indo-Slavonic model


I am not a fan of continuity theories – that much should be clear for anyone reading this blog. However, most of such proposals’ supremacist (or rather fear-of-inferiority) overtones don’t mean they have to be wrong. It just means that most of them, most of the time, most likely are.

While reading Tommenable’s comments, I thought about a potential alternative model, where one could a priori accept an identification of North Pontic cultures as ‘Indo-Slavonic’, which seems to be the Eastern European R1a continuist trend right now.

NOTE. To accept this model, one should first (not a posteriori) accept an Indo-Slavonic linguistic group on theoretical grounds, of course, and take the steppe ancestral component (and not archaeological data) as the most meaningful aspect to consider for language expansion and exchange (which we know is not the most intelligent approach to cultural or language change).

Thinking about how Genomics could challenge what mainstream Linguistics and Archaeology accepts, the only situation I can think of (using simplistic phylogeography) regarding late Khvalynsk-Sredni Stog contacts (until ca. 3300 BC) is:

  1. That the community of R1b-L51 lineages was in fact an isolated group , and not a western one – i.e. to the east within the Volga-Ural groups, or maybe to the south within the North Caucasian groups .
  2. That the R1b-Z2103 community was a huge one dominating over much of the steppe, from the Dnieper area to the Volga-Ural region (where we know they were).
  3. That R1a-M417 subclades (and especially subclade R1a-Z645) with steppe ancestry, as found in Corded Ware migrants,were only found in the North Pontic area (i.e. in Sredni Stog) during the fourth millennium (until at least 3300 BC, when Yamna substitutes it), and did not form other communities in the forest-steppe or Forest Zone (from where Corded Ware eventually expanded), as it is quite likely.
  4. That both the R1b-Z2103 and R1a-Z645 communities shared obvious genetic connections (whatever they were) around the Dnieper, that could justify a common, shared language.
Diachronic map of Eneolithic migrations in eastern Europe ca. 4000-3100 BC

Only then, if a widespread Graeco-Aryan-speaking community happened to be spread from west to east in the Pontic-Caspian steppe, with close contacts with North Pontic cultures, and having an isolated Northern Late PIE community somewhere different than West Yamna, it could leave for me a reasonable doubt of a cultural connection (maybe “Indo-Slavonic” in nature) of the North Pontic steppe. But then we would probably be stuck – yet again – with some sort of cultural diffusion event, impossible to demonstrate.

Since it is known (in Linguistics, and also in Y-DNA lineages, due to the early expansion of Z2103 subclades) that Graeco-Aryan groups separated early, this model would not be impossible.

Also a priori in favour of that model would be the early expansion of a (Northern IE-speaking) Pre-Tocharian population to the east. On the other hand, from an archaeological point of view, the group reaching Afanasevo seems to have expanded from Repin, just like the community expanding Yamna to the west of the Dnieper.

I really doubt there can be any serious discussion though, apart from amateur geneticists with a personal interest on this, because:

  • Graeco-Aryan is a Late PIE dialect, and Late PIE guesstimates are more recent than that.
  • Dialectal separation within a Late Proto-Indo-European language must have happened late, gradually, and in close contact, allowing for common innovations to spread through dialectal groups.
  • It does not make sense in terms of prehistoric cultures, since there is no direct connection or migration among steppe cultures but for the Novodanilovka and the Yamna expansions.
  • Indo-Slavonic is only supported by a handful of linguists, and not in the way or timing described in this model.

NOTE. You can read Kortlandt’s works in (also on his personal website) if you are really interested in knowing more about an Indo-Slavonic proposal, from an expert Balticist and Slavicist. However, if your intent is to demonstrate some ancient ethnic link of “your” people (whatever that means) to mythical Proto-Indo-Europeans, you would not need actual knowledge or sound theories to do that, so you can skip that part. Also, Kortlandt would probably support a later model of Indo-Slavonic expansion in the steppe, related to East Yamna, and later Sintashta, Srubna, etc…

Migration Yamna -> Corded Ware -> Bell Beaker as claimed by articles published in Nature (2015). From materials of the UAB.

If you think about it, if most modern Slavs were mainly of R1b-L23 lineages instead of R1a-Z645 (a replacement which, as it is clear know, is the consequence of a simple resurge of previous lineages in East-Central Europe, coupled with a later gradual replacement through founder effects, so no big migration history here), and Finnic speakers were mainly of R1a-Z645 lineages (whose replacement by N1c lineages seems also the consequence of quite late consecutive founder effects), I doubt we would be having this reticence to accept sound anthropological models.

So, we are speculating here for the sake of an unnecessary, naïve compromise…Just hoping to find some common ground to move on, now that the picture is clearer for everyone.

NOTE. The change of narratives where certain languages must have accompanied R1a-Z645 and N1c lineages, but in alternative ways not previously described, is obviously unjustified, if linguistic and archaeological data tell a different story. As unjustified as it is to change Yamna for “Neolithic Steppe” as homeland of Late Indo-European, to fit it with the steppe ancestry concept

See also:

Population substructure in Iberia, highest in the north-west territory (to appear in Nature)

A manuscript co-authored by Angel Carracedo, from the University of Santiago de Compostela, and (always according to him) pre-accepted in Nature, will offer more insight into the population substructure of Spain, based on autosomal DNA.

Carracedo’s lecture about DNA (in Galician), including his summary of the paper (from december 2017):

Some of the points made in the video:

  • The study shows a situation parallelling – as expected – the expansion of Spanish Medieval kingdoms during the Reconquista (and subsequent repopulation).
  • In it, the biggest surprise seems to be the greater substructure found in Galicia, the north-western Spanish territory – greater even than expected by the authors.
  • As a side note, Galicia shows a great influence from Moorish” ancestral components, due mainly to the influx from Portugal, which shows more.

It is difficult to judge only from the image and his words, but one could say that there are:

  • Certain quite old ancestral Galician groups;
    • then two – also quite old – ancestral Basque groups;
      • then more recent Galician groups;
        • and then a common, central Spanish group – including
          • a wider Asturian-Catalan group, with a western Asturian-Leonese, and an eastern Catalan subgroup;
          • and a central Castillian-Aragonese group, also with a western Castillian, and an eastern Aragonese subgroup.
Spain’s population substructure, from the video.

We thought that certain parts of the British Isles could show ancestral components related to the old population, although this has not proven exactly right, due to more recent population expansions.

However, this paper might shed light to the controversy surrounding Lusitanian (possibly Gallaico-Lusitanian) as a Pre-Celtic Indo-European group of Iberia, either slightly older as an Italo-Celtic dialect, or potentially from the Bell Beaker expansion, whose genetic imprint might have survived the Roman conquest, which apparently didn’t replace its ancestral population.

Given the presence of a central Spanish group opposed to the other minor groups – and knowing that (at least part of) the Medieval kingdoms should be related to the Occitan region – due to the Celtic expansion, and also potentially later during the Visigothic Kingdom, and the Carolingian Empire – , we can only guess that the other (north-western and Basque) groups are potentially quite old, and reflect prehistoric population structures.

Just speculating here, of course. Another interesting genetic paper to await…

Seen first in the Facebook group Iberia ADN.


Consequences of O&M 2018 (I): The latest West Yamna “outlier”


This is the first 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).

As expected, the first Y-DNA haplogroup of a sample from the North Pontic region (apart from an indigenous European I2 subclade) during its domination by the Yamna culture is of haplogroup R1b-L23, and it is dated ca. 2890-2696 BC. More specifically, it is of Z2103 subclade, the main lineage found to date in Yamna samples. The site in question is Dereivka, “in the southern part of the middle Dnieper, at the boundary between the forest-steppe and the steppe zones”.

NOTE: A bit of history for those lost here, which appear to be many: the classical Yamna culture – from previous late Khvalynsk, and (probably) Repin groupsspread west of the Don ca. 3300 BC creating a cultural-historical community – and also an early offshoot into Asia – , with mass migrations following some centuries later along the Danube to the Carpathian Basin, but also south into the Balkans, and north along the Prut. There is thus a very short time frame to find Yamna peoples shaping these massive migrations – the most likely speakers of Late Proto-Indo-European dialects – in Ukraine, compared to their most stable historical settlements east of the Don River.

There is no data on this individual in the supplementary material – since Eneolithic Dereivka samples come from stored dental remains – , but the radiocarbon date (if correct) is unequivocal: the Yamna cultural-historical community dominated over that region at that precise time. Why would the authors name it just “Ukraine_Eneolithic”? They surely took the assessment of archaeologists, and there is no data on it, so I agree this is the safest name to use for a serious paper. This would not be the first sample apparently too early for a certain culture (e.g. Catacomb in this case) which ends up being nevertheless classified as such. And it is also not impossible that it represents another close Ukraine Eneolithic culture, since ancestral cultural groups did not have borders…

NOTE. Why, on the other hand, was the sample from Zvejnieki – classified as of Latvia_LN – assumed to correspond to “Corded Ware” (like the recent samples from Plinkaigalis242 or Gyvakarai1), when we don’t have data on their cultures either? No conspiracy here, just taking assessments from different archaeologists in charge of these samples: those attributed to “Corded Ware” have been equally judged solely by radiocarbon date, but, combining the known archaeological signs of herding in the region arriving around this time with the old belief (similar to the “Iberia is the origin of Bell Beaker peoples” meme) that “only the Corded Ware culture signals the arrival of herding in the Baltic”. This assumption has been contested recently by Furholt, in an anthropological model that is now mainstream, upheld also by Anthony.

We already know that, out of three previous West Yamna samples, one shows Anatolian Neolithic ancestry, the so-called “Yamna outlier”. We also know that one sample from Yamna in Bulgaria also shows Anatolian Neolithic ancestry, with a distinct ‘southern’ drift, clustering closely to East Bell Beaker samples, as we can still see in Mathieson et al. (2018), see below. So, two “outliers” (relative to East Yamna samples) out of four samples… Now a new, fifth sample from Ukraine is another “outlier”, coinciding with (and possibly somehow late to be a part of) the massive migration waves into Central Europe and the Balkans predicted long ago by academics and now confirmed with Genomics.

I think there are two good explanations right now for its ancestral components and position in PCA:

Modified image from Mathieson et al. (2018), including also approximate location of groups from Mittnik et al. (2018), and group (transparent shape outlined by dots) formed by new Bell Beaker samples from Olalde et al. (2018). “Principal components analysis of ancient individuals. Points for 486 ancient individuals are projected onto principal components defined by 777 present-day west Eurasian individuals (grey points). Present-day individuals are shown.”

a) The most obvious one, that the Dnieper-Dniester territory must have been a melting pot, as I suggested, a region which historically connected steppe, forest steppe, and forest zone with the Baltic, as we have seen with early Baltic Neolithic samples (showing likely earlier admixture in the opposite direction). The Yamna population, a rapidly expanding “elite group of patrilineally-related families” (words from the famous 2015 genetic papers, not mine), whose only common genetic trait is therefore Y-DNA haplogroup R1b-L23, must have necessarily acquired other ancestral components of Eneolithic Ukraine during the migrations and settlements west of the Don River.

How many generations are needed for ancestral components and PCA clusters to change to that extent, in regions where only some patrilocal chiefs but indigenous populations remain, and the population probably admixed due to exogamy, back-migrations, and “resurge” events? Not many, obviously, as we see from the differences among the many Bell Beaker samples of R1b-L23 subclades from Olalde et al. (2018)

b) That this sample shows the first genetic sign of the precise population that contributed to the formation of the Catacomb culture. Since it is a hotly debated topic where and how this culture actually formed to gradually replace the Yamna culture in the central region of the Pontic-Caspian steppe, this sample would be a good hint of how its population came to be.

See e.g. for free articles on the Catacomb culture its article on the Encyclopedia of Indo-European Culture, Catacomb culture wagons of the Eurasian steppes, or The Warfare of the Northern Pontic Steppe – Forest-Steppe Pastoral Societies: 2750 – 2000 B.C. There are also many freely available Russian and Ukrainian papers on anthropometry (a discipline I don’t especially like) which clearly show early radiocarbon dates for different remains.

This could then be not ‘just another West Yamna outlier’, but would actually show meaningful ‘resurge’ of Neolithic Ukraine ancestry in the Catacomb culture.

It could be meaningul to derive hypotheses, in the same way that the late Central European CWC sample from Esperstedt (of R1a-M417 subclade) shows recent exogamy directly from the (now more probably eastern part of the) steppe or steppe-forest, and thus implies great mobility among distant CWC groups. Although, given the BB samples with elevated steppe ancestry and close PCA cluster from Olalde et al. (2018), it could also just mean exogamy from a near-by region, around the Carpathian Basin where Yamna migrants settled…

If this was the case, it would then potentially mean a “continuity” break in the steppe, in the region that some looked for as a Balto-Slavic homeland, and which would have been only later replaced by Srubna peoples with steppe ancestry (and probably R1a-Z93 subclades). We would then be more obviously left with only two options: a hypothetic ‘Indo-Slavonic’ North Caspian group to the east (supported by Kortlandt), or a Central-East European homeland near Únětice, as one of the offshoots from the North-West Indo-European group (supported by mainstream Indo-Europeanists).

How to know which is the case? We have to wait for more samples in the region. For the moment, the date seems too early for the known radiocarbon dating of most archaeological remains of the Catacomb Culture.

Diachronic map of Late Copper Age migrations including steppe groups ca. 2600-2250 BC

An important consequence of the addition of these “Yamna outliers” for the future of research on Indo-European migrations is that, especially if confirmed as just another West Yamna sample – with more, similar samples – , early Palaeo-Balkan peoples migrating south of the Danube and later through Anatolia may need to be judged not only in terms of ancestral components or PCA (as in the paper on Minoans and Mycenaeans), but also and more decisively using phylogeography, especially with the earliest samples potentially connected with such migrations.

NOTE. Regarding the controversy (that some R1b European autochthonous continuists want to create) over the origin of the R1b-L151 lineages, we cannot state its presence for sure in Yamna territory right now, but we already have R1b-M269 in the eastern Pontic-Caspian steppe during the Neolithic-Chalcolithic transition, then R1b-L23 and subclades (mainly R1b-Z2013, but also one xZ2103, xL51 which suggests its expansion) in the region before and during the Yamna expansion, and now we have L51 subclades with elevated steppe ancestry in early East Bell Beakers, which most likely descended from Yamna settlers in the Carpathian Basin (yet to be sampled).

Even without express confirmation of its presence in the steppe, the alternative model of a Balkan origin seems unlikely, given the almost certain continuity of expanding Yamna clans as East Bell Beaker ones, in this clearly massive and relatively quick expansion that did not leave much time for founder effects. But, of course, it is not impossible to think about a previously hidden R1b-L151 community in the Carpathian Basin yet to be discovered, adopting North-West Indo-European (by some sort of founder effect) brought there by Yamna peoples of exclusively R1b-Z2103 lineages. As it is not impossible to think about a hidden and ‘magically’ isolated community of haplogroup R1a-M417 in Yamna waiting to be discovered…Just not very likely, either option.

As to why this sample or the other Bell Beaker samples “solve” the question of R1a-Z645 subclades (typical of Corded Ware migrants) not expanding with Yamna, it’s very simple: it doesn’t. What should have settled that question – in previous papers, at least since 2015 – is the absence of this subclade in elite chiefs of clans expanded from Khvalynsk, Yamna, or their only known offshoots Afanasevo and Bell Beaker. Now we only have still more proof, and no single ‘outlier’ in that respect.

No haplogroup R1a among hundreds of samples from a regionally extensive sampling of the only cultures mainstream archaeologists had thoroughly described as potentially representing Indo-European-speakers should mean, for any reasonable person (i.e. without a personal or professional involvement in an alternative hypothesis), that Corded Ware migrants (as expected) did not stem from Yamna, and thus did not spread Late Indo-European dialects.

This haplogroup’s hegemonic presence in North-Eastern Europe – and the lack of N1c lineages until after the Bronze Age – coinciding with dates when Uralicists have guesstimated Uralic dialectal expansion accross this wide region makes the question of the language spread with CWC still clearer. The only surprise would have been to find a hidden and isolated community of R1a-Z645 lineages clearly associated with the Yamna culture.

NOTE. A funny (however predictable) consequence for R1a autochthonous continuists of Northern or Eastern European ancestry: forum commentators are judging if this sample was of the Yamna culture or spoke Indo-European based on steppe component and PCA cluster of the few eastern Yamna samples which define now (you know, with the infallible ‘Yamnaya ancestral component’) the “steppe people” who spoke the “steppe language”™ – including, of course, North-Eastern European Late Neolithic

Not that radiocarbon dates or the actual origin of this sample cannot be wrong, mind you, it just strikes me how twisted such biased reasonings may be, depending on the specific sample at hand… Denial, anger, and bargaining, including shameless circular reasoning – we know the drill: we have seen it a hundred times already, with all kinds of supremacists autochthonous continuists who still today manage to place an oudated mythical symbolism on expanding Proto-Indo-Europeans, or on regional ethnolinguistic continuity…

More detailed posts on the new samples from O&M 2018 and their consequences for the Indo-European demic diffusion to come, indeed…

See also: