Aquitanians and Iberians of haplogroup R1b are exactly like Indo-Iranians and Balto-Slavs of haplogroup R1a

eba-indo-iranian-balto-slavs

The final paper on Indo-Iranian peoples, by Narasimhan and Patterson (see preprint), is soon to be published, according to the first author’s Twitter account.

One of the interesting details of the development of Bronze Age Iberian ethnolinguistic landscape was the making of Proto-Iberian and Proto-Basque communities, which we already knew were going to show R1b-P312 lineages, a haplogroup clearly associated during the Bell Beaker period with expanding North-West Indo-Europeans:

From the Bronze Age (~2200–900 BCE), we increase the available dataset from 7 to 60 individuals and show how ancestry from the Pontic-Caspian steppe (Steppe ancestry) appeared throughout Iberia in this period, albeit with less impact in the south. The earliest evidence is in 14 individuals dated to ~2500–2000 BCE who coexisted with local people without Steppe ancestry. These groups lived in close proximity and admixed to form the Bronze Age population after 2000 BCE with ~40% ancestry from incoming groups. Y-chromosome turnover was even more pronounced, as the lineages common in Copper Age Iberia (I2, G2, and H) were almost completely replaced by one lineage, R1b-M269.

iberia-admixture-y-dna
Proportion of ancestry derived from central European Beaker/Bronze Age populations in Iberians from the Middle Neolithic to the Iron Age (table S15). Colors indicate the Y-chromosome haplogroup for each male. Red lines represent period of admixture. Modified from Olalde et al. (2019).

The arrival of East Bell Beakers speaking Indo-European languages involved, nevertheless, the survival of the two non-IE communities isolated from each other – likely stemming from south-western France and south-eastern Iberia – thanks to a long-lasting process of migration and admixture. There are some common misconceptions about ancient languages in Iberia which may have caused some wrong interpretations of the data in the paper and elsewhere:

NOTE. A simple reading of Iberian prehistory would be enough to correct these. Two recent books on this subject are Villar’s Indoeuropeos, iberos, vascos y otros parientes and Vascos, celtas e indoeuropeos. Genes y lenguas.

Iberian languages were spoken at least in the Mediterranean and the south (ca. “1/3 of Iberia“) during the Bronze Age.

Nope, we only know the approximate location of Iberian culture and inscriptions from the Late Iron Age, and they occupy the south-eastern and eastern coastal areas, but before that it is unclear where they were spoken. In fact, it seems evident now that the arrival of Urnfield groups from the north marks the arrival of Celtic-speaking peoples, as we can infer from the increase in Central European admixture, while the expansion of anthropomorphic stelae from the north-west must have marked the expansion of Lusitanian.

Vasconic was spoken in both sides of the Pyrenees, as it was in the Middle Ages.

Wrong. One of the worst mistakes I am seeing in many comments since the paper was published, although admittedly the paper goes around this problem talking about “Modern Basques”. Vasconic toponyms appear south of the Pyrenees only after the Roman conquests, and tribes of the south-western Pyrenees and Cantabrian regions were likely Celtic-speaking peoples. Aquitanians (north of the western Pyrenees) are the only known ancient Vasconic-speaking population in proto-historic times, ergo the arrival of Bell Beakers in Iberia was most likely accompanied by Indo-European languages which were later replaced by Celtic expanding from Central Europe, and Iberian expanding from south-east Iberia, and only later with Latin and Vasconic.

Ligurian is non-Indo-European, and Lusitanian is Celtic-like, so Iberia must have been mostly non-Indo-European-speaking.

The fragmentary material available on Ligurian is enough to show that phonetically it is a NWIE dialect of non-Celtic, non-Italic nature, much like Lusitanian; that is, unless you follow laryngeals up to Celtic or Italic, in which case you can argue anything about this or any other IE language, as people who reconstruct laryngeals for Baltic in the common era do.

EDIT (19 Mar 2019): It was not clear enough from this paragraph, because Ligurian-like languages in NE Iberia is just a hypothesis based on the archaeological connection of the whole southern France Bell Beaker region. My aim was to repeat the idea that Old European topo-hydronymy is older in NE Iberia (as almost anywhere in Iberia) than Iberian toponymy, so the initial hypothesis is that:

  1. a Palaeo-European language (as Villar puts it) expanded into most regions of Iberia in ancient times (he considered at some point the Mesolithic, but that is obviously wrong, as we know now); then
  2. Celts expanded at least to the Ebro River Basin; then
  3. Iberians expanded to the north and replaced these in NE Iberia; and only then
  4. after the Roman invasion, around the start of the Common Era, appear Vasconic toponyms south of the Pyrenees.

Lusitanian obviously does not qualify as Celtic, lacking the most essential traits that define Celticness…Unless you define “(Para-)Celtic” as Pre-Proto-Celtic-like, or anything of the sort to support some Atlantic continuity, in which case you can also argue that Pre-Italic or Pre-Germanic are Celtic, because you would be essentially describing North-West Indo-European

If Basques have R1b, it’s because of a culture of “matrilocality” as opposed to the “patrilocality” of Indo-Europeans

So wrong it hurts my eyes every time I read this. Not only does matrilocality in a regional group have few known effects in genetics, but there are many well-documented cases of population replacement (with either ancestry or Y-DNA haplogroups, or both) without language replacement, without a need to resort to “matrilineality” or “matrilocality” or any other cultural difference in any of these cases.

In fact, it seems quite likely now that isolated ancient peoples north of the Pyrenees will show a gradual replacement of surviving I2a lineages by neighbouring R1b, while early Iberian R1b-DF27 lineages are associated with Lusitanians, and later incoming R1b-DF27 lineages (apart from other haplogroups) are most likely associated with incoming Celts, which must have remained in north-central and central-east European groups.

NOTE. Notice how R1a is fully absent from all known early Indo-European peoples to date, whether Iberian IE, British IE, Italic, or Greek. The absence of R1a in Iberia after the arrival of Celts is even more telling of the origin of expanding Celts in Central Europe.

I haven’t had enough time to add Iberian samples to my spreadsheet, and hence neither to the ASoSaH texts nor maps/PCAs (and I don’t plan to, because it’s more efficient for me to add both, Asian and Iberian samples, at the same time), but luckily Maciamo has summed it up on Eupedia. Or, graphically depicted in the paper for the southeast:

iberia-haplogroups
Y chromosome haplogroup composition of individuals from southeast Iberia during the past 2000 years. The general Iberian Bronze and Iron Age population is included for comparison. Modified from Olalde et al. (2019).

Does this continued influx of Y-DNA haplogroups in Iberia with different cultures represent permanent changes in language? Are, therefore, modern Iberian languages derived from Lusitanian, Sorothaptic/Celtic, Greek, Phoenician, East or West Germanic, Hebrew, Berber, or Arabic languages? Obviously not. Same with Italy (see the recent preprint on modern Italians by Raveane et al. 2018), with France, with Germany, or with Greece.

If that happens in European regions with a known ancient history, why would the recent expansions and bottlenecks of R1b in modern Basques (or N1c around the Baltic, or R1a in Slavs) in the Middle Ages represent an ancestral language surviving into modern times?

Indo-Iranians

If something is clear from Narasimhan, Patterson, et al. (2018), is that we know finally the timing of the introduction and expansion of R1a-Z645 lineages among Indo-Iranians.

We could already propose since 2015 that a slow admixture happened in the steppes, based on archaeological finds, due to settlement elites dominating over common peoples, coupled with the known Uralic linguistic traits of Indo-Iranian (and known Indo-Iranian influence on Finno-Ugric) – as I did in the first version of the Indo-European demic diffusion model.

The new huge sampling of Sintashta – combined with that of Catacomb, Poltavka, Potapovka, Andronovo, and Srubna – shows quite clearly how this long-term admixture process between Uralic peoples and Indo-Iranians happened between forest-steppe CWC (mainly Abashevo) and steppe groups. The situation is not different from that of Iberia ca. 2500-2000 BC; from Narasimhan, Patterson, et al. (2018):

We combined the newly reported data from Kamennyi Ambar 5 with previously reported data from the Sintashta 5 individuals (10). We observed a main cluster of Sintashta individuals that was similar to Srubnaya, Potapovka, and Andronovo in being well modeled as a mixture of Yamnaya-related and Anatolian Neolithic (European agriculturalist-related) ancestry.

Even with such few words referring to one of the most important data in the paper about what happened in the steppes, Wang et al. (2018) help us understand what really happened with this simplistic concept of “steppe ancestry” regarding Yamna vs. Corded Ware differences:

anatolia-neolithic-steppe-eneolithic
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).”

As with Iberia (or any prehistoric region), the details of how exactly this language change happened are not evident, but we only need a plausible explanation coupled with archaeology and linguistics. Poltavka, Potapovka, and Sintashta samples – like the few available Iberian ones ca. 2500-2000 BC – offer a good picture of the cohabitation of R1b-L23 (mainly Z2103) and R1a-Z645 (mainly Z93+): a glimpse at the likely presence of R1a-Z93 within settlements – which must have evolved as the dominant elites – in a society where the majority of the population was initially formed by nomad herders (probably most R1b-Z2103), who were usually buried outside of the main settlements.

Will the upcoming Narasimhan, Patterson et al. (2019) deal with this problem of how R1a-M417 replaced R1b-M269, and how the so-called “Steppe_MLBA” (i.e. Corded Ware) ancestry admixed with “Steppe_EMBA” (i.e. Yamnaya) ancestry in the steppes, and which one of their languages survived in the region (that is, the same the Reich Lab has done with Iberia)? Not likely. The ‘genetic wars’ in Iberia deal with haplogroup R1b-P312, and how it was neither ‘native’ nor associated with Basques and non-Indo-European peoples in general. The ‘genetic wars’ in South Asia are concerned with the steppe origin of R1a, to prove that it is not a ‘native’ haplogroup to India, and thus neither are Indo-Aryan languages. To each region a politically correct account of genetic finds, with enough care not to fully dismiss national myths, it seems.

NOTE. Funnily enough, these ‘genetic wars’ are the making of geneticists since the 1990s and 2000s, so we are still in the midst of mostly internal wars caused by what they write. Just as genetic papers of the 2020s will most likely be a reaction to what they are writing right now about “steppe ancestry” and R1a. You won’t find much change to the linguistic reconstruction in this whole period, except for the most multicolored glottochronological proposals…

The first author of the paper has engaged, as far as I could see in Twitter, in dialogue with Hindu nationalists who try to dismiss the arrival of steppe ancestry and R1a into South Asia as inconclusive (to support the potential origin of Sanskrit millennia ago in the Indus Valley Civilization). How can geneticists deal with the real problem here (the original ethnolinguistic group expanding with Corded Ware), when they have to fend off anti-steppists from Europe and Asia? How can they do it, when they themselves are part of the same societies that demand a politically correct presentation of data?

This is how the data on the most likely Indo-Iranian-speaking region should be presented in an ideal world, where – as in the Iberia paper – geneticists would look closely to the Volga-Ural region to discover what happened with Proto-Indo-Iranians from their earliest to their latest stage, instead of constantly looking for sites close to the Indus Valley to demonstrate who knows what about modern Indian culture:

indo-iranian-admixture-similar-iberians
Tentative map of the Late PIE and Indo-Iranian community in the Volga-Ural steppes since the Eneolithic. Proportion of ancestry derived from central European Corded Ware peoples. Colors indicate the Y-chromosome haplogroup for each male. Red lines represent period of admixture. Modified from Olalde et al. (2019).

Now try and tell Hindu nationalists that Sanskrit expanded from an Early Bronze Age steppe community of R1b-rich nomadic herders that spoke Pre-Indo-Iranian, which was dominated and eventually (genetically) mostly replaced by elite Uralic-speaking R1a peoples from the Russian forest, hence the known phonetic (and some morphological) traits that remained. Good luck with the Europhobic shitstorm ahead..

Balto-Slavic

Iberian cultures, already with a majority of R1b lineages, show a clear northward expansion over previously Urnfield-like groups of north-east Iberia and Mediterranean France (which we now know probably represent the migration of Celts from central Europe). Similarly, Eastern Balts already under a majority of R1a lineages expanded likely into the Baltic region at the same time as the outlier from Turlojiškė (ca. 1075 BC), which represents the first obvious contacts of central-east Europe with the Baltic.

Iberia shows a more recent influx of central and eastern Mediterranean peoples, one of which eventually succeeded in imposing their language in Western Europe: Romans were possibly associated mainly with R1b-U152, apart from many other lineages. Proto-Slavs probably expanded later than Celts, too, connected to the disintegration of the Lusatian culture, and they were at some point associated with R1a-M458 and R1a-Z280(xZ92) lineages, apart from others already found in Early Slavs.

pca-balto-slavs-tollense-valley
PCA of central-eastern European groups which may have formed the Balto-Slavic-speaking community derived from Bell Beaker, evident from the position ‘westwards’ of CWC in the PCA, and surrounding cultures. Left: Early Bronze Age. Right: Tollense Valley samples.

This parallel between Iberia and eastern Europe is no coincidence: as Europe entered the Bronze Age, chiefdom-based systems became common, and thus the connection of ancestry or haplogroups with ethnolinguistic groups became weaker.

What happened earlier (and who may represent the Pre-Balto-Slavic community) will be clearer when we have enough eastern European samples, but basically we will be able to depict this admixture of NWIE-speaking BBC-derived peoples with Uralic-speaking CWC-derived groups (since Uralic is known to have strongly influenced Balto-Slavic), similar to the admixture found in Indo-Iranians, more or less like this:

iberian-admixture-balto-slavic
Tentative map of the North-West Indo-European and Balto-Slavic community in central-eastern Europe since the East Bell Beaker expansion. Proportion of ancestry derived from Corded Ware peoples. Colors indicate the Y-chromosome haplogroup for each male. Red lines represent period of admixture. Modified from Olalde et al. (2019).

The Early Scythian period marked a still stronger chiefdom-based system which promoted the creation of alliances and federation-like groups, with an earlier representation of the system expanding from north-eastern Europe around the Baltic Sea, precisely during the spread of Akozino warrior-traders (in turn related to the Scythian influence in the forest-steppes), who are the most likely ancestors of most N1c-V29 lineages among modern Germanic, Balto-Slavic, and Volga-Finnic peoples.

Modern haplogroup+language = ancient ones?

It is not difficult to realize, then, that the complex modern genetic picture in Eastern Europe and around the Urals, and also in South Asia (like that of the Aegean or Anatolia) is similar to the Iron Age / medieval Iberian one, and that following modern R1a as an Indo-European marker just because some modern Indo-European-speaking groups showed it was always a flawed methodology; as flawed as following R1b for ancient Vasconic groups, or N1c for ancient Uralic groups.

Why people would argue that haplogroups mean continuity (e.g. R1b with Basques, N1c with Finns, R1a with Slavs, etc.) may be understood, if one lives still in the 2000s. Just like why one would argue that Corded Ware is Indo-European, because of Gimbutas’ huge influence since the 1960s with her myth of “Kurgan peoples”. Not many denied these haplogroup associations, because there was no reason to do it, and those who did usually aligned with a defense of descriptive archaeology.

However, it is a growing paradox that some people interested in genetics today would now, after the Iberian paper, need to:

  • accept that ancient Iberians and probably Aquitanians (each from different regions, and probably from different “Basque-Iberian dialects” in the Chalcolithic, if both were actually related) show eventually expansions with R1b-L23, the haplogroup most obviously associated with expanding Indo-Europeans;
  • acknowledge that modern Iberians have many different lineages derived from prehistoric or historic peoples (Celts, Phoenicians, Greeks, Romans, Jews, Goths, Berbers, Arabs), which have undergone different bottlenecks, the last ones during the Reconquista, but none of their languages have survived;
  • realize that a similar picture is to be found everywhere in central and western Europe since the first proto-historic records, with language replacement in spite of genetic continuity, such as the British Isles (and R1b-L21 continuity) after the arrival of Celts, Romans, Anglo-Saxons, Vikings, or Normans;
  • but, at the same time, continue blindly asserting that haplogroup R1a + “steppe ancestry” represent some kind of supernatural combination which must show continuity with their modern Indo-Iranian or Balto-Slavic language from time immemorial.
sintashta-y-dna
Replacement of R1b-L23 lineages during the Early Bronze Age in eastern Europe and in the Eurasian steppes: emergence of R1a in previous Yamnaya and Bell Beaker territories. Modified from EBA Y-DNA map.

Behave, pretty please

The ‘conservative’ message espoused by some geneticists and amateur genealogists here is basically as follows:

  • Let’s not rush to new theories that contradict the 2000s, lest some people get offended by granddaddy not being these pure whatever wherever as they believed, and let’s wait some 5, 10, or 20 years, as long as necessary – to see if some corner of the Yamna culture shows R1a, or some region in north-eastern Europe shows N1c, or some Atlantic Chalcolithic sample shows R1b – to challenge our preferred theories, if we actually need to challenge anything at all, because it hurts too much.
  • Just don’t let many of these genetic genealogists or academics of our time be unhappy, pretty please with sugar on top, and let them slowly adapt to reality with more and more pet theories to fit everything together (past theories + present data), so maybe when all of them are gone, within 50 or 70 years, society can smoothly begin to move on and propose something closer to reality, but always as politically correct as possible for the next generations.
  • For starters, let’s discuss now (yet again) that Bell Beakers may not have been Indo-European at all, despite showing (unlike Corded Ware) clearly Yamna male lineages and ancestry, because then Corded Ware and R1a could not have been Indo-European and that’s terrible, so maybe Bell Beakers are too brachycephalic to speak Indo-European or something, or they were stopped by the Fearsome Tisza River, or they are not pure Dutch Single Grave in The South hence not Indo-European, or whatever, and that’s why Iron Age Iberians or Etruscans show non-Indo-European languages. That’s not disrespectful to the history of certain peoples, of course not, but talking about the evident R1a-Uralic connection is, because this is The South, not The North, and respect works differently there.
  • Just don’t talk about how Slavs and Balts enter history more than 1,500 years later than Indo-European peoples in Western and Southern Europe, including Iberia, and assume a heroic continuity of Balts and Slavs as pure R1a ‘steppe-like’ peoples dominating over thousands of kms. in the Baltic, Fennoscandia, eastern Europe, and northern Asia for 5,000 years, with multiple Balto-Slavs-over-Balto-Slavs migrations, because these absolute units of Indo-European peoples were a trip and a half. They are the Asterix and Obelix of white Indo-European prehistory.
  • Perhaps in the meantime we can also invent some new glottochronological dialectal scheme that fits the expansion of Sredni Stog/Corded Ware with (Germano-?)Indo-Slavonic separated earlier than any other Late PIE dialect; and Finno-Volgaic later than any other Uralic dialect, in the Middle Ages, with N1c.
balto-slavic-pca
Genetic structure of the Balto-Slavic populations within a European context according to the three genetic systems, from Kushniarevich et al. (2015). Pure Balto-Slavs from…hmm…yeah this…ancient…region…or people…cluster…Whatever, very very steppe-like peoples, the True Indo-Europeans™, so close to Yamna…almost as close as Finno-Ugrians.

To sum up: Iberia, Italy, France, the British Isles, central Europe, the Balkans, the Aegean, or Anatolia, all these territories can have a complex history of periodic admixture and language replacement everywhere, but some peoples appearing later than all others in the historical record (viz. Basques or Slavs) apparently cannot, because that would be shameful for their national or ethnic myths, and these should be respected.

Ignorance of the own past as a blank canvas to be filled in with stupid ethnolinguistic continuity, turned into something valuable that should not be challenged. Ethnonationalist-like reasoning proper of the 19th century. How can our times be called ‘modern’ when this kind of magical thinking is still prevalent, even among supposedly well-educated people?

Related

Haplogroup R1b-M167/SRY2627 linked to Celts expanding with the Urnfield culture

bronze-age-late-urnfield

As you can see from my interest in the recently published Olalde et al. (2019) Iberia paper, once you accept that East Bell Beakers expanded North-West Indo-European, the most important question becomes how did its known dialects spread to their known historic areas.

We already had a good idea about the expansion of Celts, based on proto-historical accounts, fragmentary languages, and linguistic guesstimates, but the connection of Celtic with either Urnfield or slightly later Hallstatt/La Tène was always blurred, due to the lack of precise data on population movements.

The latest paper on Iberia is interesting for many details, such as:

  • The express dismissal of the newest pet theory based on the simplistic “steppe ancestry = IE”: the obsessive comparisons of Dutch Bell Beakers as the origin of basically anything that moves in Europe.
  • A discrete influx of North African ancestry in certain samples before the Moorish invasion (which was probably mediated by peoples of North African rather than Levantine admixture).
  • The finding of very Mycenaean-like Greek colonies of the 5th century (interestingly, under R1b lineages).
iberia-celts-romans
Modified from section of PCA of ancient samples by Olalde et al. (2019). “IE Iberia” refers to Pre-Celtic Indo-European languages of Iberia, such as Galaico-Lusitanian in the west (see more on Lusitanian), and a potentially Ligurian-related language in the North-East and southern France.

The paper is, however, of particular importance from the perspective of historical linguistics. It confirms that:

  • Celtic-speaking peoples expanded in Iberia likely during the Late Bronze Age – Early Iron Age (probably with the Urnfield culture, before 1000 BC) with North/Central European ancestry.

NOTE. The paper marks what are believed to be the boundaries of non-Indo-European languages during the Iron Age in later times, extrapolating that situation to the past. Mediterranean sites with Iberian traits (ca. 6th century on) were probably non-Indo-European-speaking tribes, but it is unclear what happened in the centuries before their sampling, and there are no clear boundaries. These incoming Celts from central Europe with the Urnfield culture makes it very likely that the Iberian expansion to the north happened later, incorporating thus this central European ancestry in the process. The southern (orientalizing, Tartessian) site of La Angorrilla shows incineration and influence from Phoenician settlers, and their actual language is also far from clear. The other investigated samples, with higher central European contribution, are from Celtiberian sites.

  • The slightly later arrival of (Phoenician, Greek and) Latin-speaking peoples into Iberia is marked by Central/Eastern Mediterranean and North African ancestry.
iberia-migrations-celts-romans
Expansion of different ancestry components in Iberia during Prehistory. Modified from Olalde et al. (2019) to include labels with populations expanding with each component.

While both confirm what was more or less already known about the oldest attested NWIE dialects, and further support the role of East Bell Beakers in expanding North-West Indo-European, the first part is interesting for two main reasons:

  1. Koch’s Celtic from the West hypothesis, which made a recent comeback with a renewed model based on “steppe ancestry”, is once again rejected in population genomics, as expected. At this point I doubt this will mean anything to the supporters of the theory (because you can propose as many “Celtic-over-Celtic” layers as you want), but if you are not obsessed with autochthonous continuity of Celtic languages in the Atlantic area we might begin to judge the most correct dialectal split (and thus classification) among those proposed to date, based on ancestry and haplogroup expansions.
  2. We believed in the 2000s that the expansion of haplogroup R1b-M167 (TMRCA ca. 1100 BC for YTree or 1700 BC for YFull) was coupled with the expansion of Iberians from the Pyrenees, in turn (thus) closely related to Basques. This non-IE presence has been contested with toponymic data in linguistics, and with the testing of many modern samples and the subsequent discovery of the widespread distribution of the subclade in western and northern Europe. Now it has become even more likely (lacking confirmation with aDNA) that this haplogroup expanded with Celts.

NOTE. Regarding R1b SNPs, YTree has more samples (and thus more SNPs) to work with estimates, due to its connection with FTDNA groups, so it is in principle more reliable (although estimates were calculated in 2017). Nevertheless, the methods to estimate the age of the MRCA are different between YTree and YFull.

df27-m167-z262-mcdonald
YTree estimations of TMRCA for R1b-Z262 (left) and R1b-M167 (right).

Why this is important has to do with the realization that Celts must have expanded explosively in all directions during the estimated range for Common Celtic (ca. 1500-1000 BC), and as such R1b-M167 is probably going to be one of the clear Y-DNA markers of the Celtic expansion, when it appears in the ancient DNA record, maybe in new SNP calls from samples of the Olalde et al. (2019) paper, or in future Urnfield/Hallstatt/La Tène papers.

Sister clades derived from R1b-Z262 (TMRCA ca. 1650 BC for YTree, or 2700 for YFull), although sharing a quite old origin, may have taken part in the same communities that expanded R1b-M167, likely from some point in central Europe, possibly as remnants of a previous (Tumulus culture?) central European expansion, as the sample SZ5 from Szólád (R1b-CTS1595) and the distribution of modern samples suggest.

r1b-df27-m167-sry2627
Left: Modern distribution of upstream clade L176.2 (YFull R1b-CTS4188); Right: Modern distribution of M167. Both include later expansions within Iberia (probably with the Crown of Aragon during the Reconquista). Contour maps of the derived allele frequencies of the SNPs analyzed in Solé-Morata et al. (2017).

The Celtic expansion might not have been a mass migration of peoples replacing all male lines of their controlled territories (as was common in the Neolithic and Chalcolithic), because of the Bronze Age dominant chiefdom-based system that relied on alliances, but it is becoming clear that Early Celts are also going to show the expansion of certain successful male lineages.

Oh, and you can say goodbye to the autochthonous “Vasconic = R1b-DF27” (latest heir of the “Vasconic = R1b-P312”) theory, too, if – for some strange reason – you hadn’t already.

EDIT (16 MAR) Just in case the wording is not clear: the fact that this haplogroup most likely expanded with Celts does not mean that its lineages didn’t become eventually incorporated into Iberian cultures and adopted non-IE languages: some of them probably did at some point, in some regions of northern Iberia, and most were certainly later incorporated to the Roman civilization and spoke Latin, then to the medieval kingdoms with their languages, and so on until the present day… Only those eventually associated with Iron Age Aquitanians may have retained their non-IE language, unless those lineages today associated with Basques were incorporated later to the Basque-speaking regions by expanding medieval kingdoms. A complex picture repeated everywhere in Europe: no haplogroup+language continuity in sight, anywhere.

NOTE: This here is currently the most likely interpretation of data based on estimations of mutations; it is not confirmed with ancient samples.

Related

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

bronze_age_iberia

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.

iberia-y-dna-mtdna
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.

iberia-steppe-admixture
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.

bronze-age-tollense
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 & 26 MAR 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.

Related:

Migrations painted by Irish and Scottish genetic clusters, and their relationship with British and European ones

ireland-britain-cluster

Interesting and related publications, now appearing in pairs…

1. The Irish DNA Atlas: Revealing Fine-Scale Population Structure and History within Ireland, by Gilbert et al., in Scientific Reports (2017).

Abstract:

The extent of population structure within Ireland is largely unknown, as is the impact of historical migrations. Here we illustrate fine-scale genetic structure across Ireland that follows geographic boundaries and present evidence of admixture events into Ireland. Utilising the ‘Irish DNA Atlas’, a cohort (n = 194) of Irish individuals with four generations of ancestry linked to specific regions in Ireland, in combination with 2,039 individuals from the Peoples of the British Isles dataset, we show that the Irish population can be divided in 10 distinct geographically stratified genetic clusters; seven of ‘Gaelic’ Irish ancestry, and three of shared Irish-British ancestry. In addition we observe a major genetic barrier to the north of Ireland in Ulster. Using a reference of 6,760 European individuals and two ancient Irish genomes, we demonstrate high levels of North-West French-like and West Norwegian-like ancestry within Ireland. We show that that our ‘Gaelic’ Irish clusters present homogenous levels of ancient Irish ancestries. We additionally detect admixture events that provide evidence of Norse-Viking gene flow into Ireland, and reflect the Ulster Plantations. Our work informs both on Irish history, as well as the study of Mendelian and complex disease genetics involving populations of Irish ancestry.

european-ancestry-british-isles
The European ancestry profiles of 30 Irish and British clusters. (a) The total ancestry contribution summarised by majority European country of origin to each of the 30 Irish and British clusters. (b) (left) The ancestry contributions of 19 European clusters that donate at least 2.5% ancestry to any one Irish or British cluster. (right) The geographic distribution of the 19 European clusters, shown as the proportion of individuals in each European region belonging to each of the 19 European clusters. The proportion of individuals form each European region not a member of the 19 European clusters is shown in grey. Total numbers of individuals from each region are shown in white text. Not all Europeans included in the analysis were phenotyped geographically. The figure was generated in the statistical software language R46, version 3.4.1, using various packages. The map of Europe was sourced from the R software package “mapdata” (https://CRAN.R-project.org/package=mapdata).

2. New preprint on BioRxiv, Insular Celtic population structure and genomic footprints of migration, by Byrne, Martiniano et al. (2017).

Abstract:

Previous studies of the genetic landscape of Ireland have suggested homogeneity, with population substructure undetectable using single-marker methods. Here we have harnessed the haplotype-based method fineSTRUCTURE in an Irish genome-wide SNP dataset, identifying 23 discrete genetic clusters which segregate with geographical provenance. Cluster diversity is pronounced in the west of Ireland but reduced in the east where older structure has been eroded by historical migrations. Accordingly, when populations from the neighbouring island of Britain are included, a west-east cline of Celtic-British ancestry is revealed along with a particularly striking correlation between haplotypes and geography across both islands. A strong relationship is revealed between subsets of Northern Irish and Scottish populations, where discordant genetic and geographic affinities reflect major migrations in recent centuries. Additionally, Irish genetic proximity of all Scottish samples likely reflects older strata of communication across the narrowest inter-island crossing. Using GLOBETROTTER we detected Irish admixture signals from Britain and Europe and estimated dates for events consistent with the historical migrations of the Norse-Vikings, the Anglo-Normans and the British Plantations. The influence of the former is greater than previously estimated from Y chromosome haplotypes. In all, we paint a new picture of the genetic landscape of Ireland, revealing structure which should be considered in the design of studies examining rare genetic variation and its association with traits.

Here are some interesting excerpts (emphasis mine):

Population structure in Ireland

The geographical distribution of this deep subdivision of Leinster resembles pre-Norman territorial boundaries which divided Ireland into fifths (cúige), with north Leinster a kingdom of its own known as Meath (Mide) [15]. However interpreted, the firm implication of the observed clustering is that despite its previously reported homogeneity, the modern Irish population exhibits genetic structure that is subtly but detectably affected by ancestral population structure conferred by geographical distance and, possibly, ancestral social structure.

ChromoPainter PC1 demonstrated high diversity amongst clusters from the west coast, which may be attributed to longstanding residual ancient (possibly Celtic) structure in regions largely unaffected by historical migration. Alternatively, genetic clusters may also have diverged as a consequence of differential influence from outside populations. This diversity between western genetic clusters cannot be explained in terms of geographic distance alone.

In contrast to the west of Ireland, eastern individuals exhibited relative homogeneity; (…) The overall pattern of western diversity and eastern homogeneity in Ireland may be explained by increased gene flow and migration into and across the east coast of Ireland from geographically proximal regions, the closest of which is the neighbouring island of Britain.

Analysis of variance of the British admixture component in cluster groups showed a significant difference (p < 2×10-16), indicating a role for British Anglo-Saxon admixture in distinguishing clusters, and ChromoPainter PC2 was correlated with the British component (p < 2×10-16), explaining approximately 43% of the variance. PC2 therefore captures an east to west Anglo-Celtic cline in Irish ancestry. This may explain the relative eastern homogeneity observed in Ireland, which could be a result of the greater English influence in Leinster and the Pale during the period of British rule in Ireland following the Norman invasion, or simply geographic proximity of the Irish east coast to Britain. Notably, the Ulster cluster group harboured an exceptionally large proportion of the British component (Fig 1D and 1E), undoubtedly reflecting the strong influence of the Ulster Plantations in the 17th century and its residual effect on the ethnically British population that has remained.

ireland-population-structure
Fine-grained population structure in Ireland. (A) fineSTRUCTURE clustering dendrogram for 1,035 Irish individuals. Twenty-three clusters are defined, which are combined into cluster groups for clusters that are neighbouring in the dendrogram, overlapping in principal component space (B) and sampled from regions that are geographically contiguous. Details for each cluster in the dendrogram are provided in S1 Fig. (B) Principal components analysis (PCA) of haplotypic similarity, based on ChromoPainter coancestry matrix for Irish individuals. Points are coloured according to cluster groups defined in (A); the median location of each cluster group is plotted. (C) Map of Irelandshowing the sampling location for a subset of 588 individuals analysed in (A) and (B), coloured by cluster group. Points have been randomly jittered within a radius of 5 km to preserve anonymity. Precise sampling location for 44 Northern Irish individuals from the People of the British Isles dataset was unknown; these individuals are plotted geometrically in a circle. (D) “British admixture component” (ADMIXTURE estimates; k=2) for Irish cluster groups. This component has the largest contribution in ancient Anglo-Saxons and the SEE cluster. (E) Linear regression of principal component 2 (B) versus British admixture component (r2 = 0.43; p < 2×10-16). Points are coloured by cluster group. (Standard error for ADMIXTURE point estimates presented in S11 Fig.)

On the genetic structure of the British Isles

The genetic substructure observed in Ireland is consistent with long term geographic diversification of Celtic populations and the continuity shown between modern and Early Bronze Age Irish people

Clusters representing Celtic populations harbouring less Anglo-Saxon influence separate out above and below SEE on PC4. Notably, northern Irish clusters (NLU), Scottish (NISC, SSC and NSC), Cumbria (CUM) and North Wales (NWA) all separate out at a mutually similar level, representing northern Celtic populations. The southern Celtic populations Cornwall (COR), south Wales (SWA) and south Munster (SMN) also separate out on similar levels, indicating some shared haplotypic variation between geographically proximate Celtic populations across both Islands. It is notable that after the split of the ancestrally divergent Orkney, successive ChromoPainter PCs describe diversity in British populations where “Anglo-saxonization” was repelled [22]. PC3 is dominated by Welsh variation, while PC4 in turn splits North and South Wales significantly, placing south Wales adjacent to Cornwall and north Wales at the other extreme with Cumbria, all enclaves where Brittonic languages persisted.

In an interesting symmetry, many Northern Irish samples clustered strongly with southern Scottish and northern English samples, defining the Northern Irish/Cumbrian/Scottish (NICS) cluster group. More generally, by modelling Irish genomes as a linear mixture of haplotypes from British clusters, we found that Scottish and northern English samples donated more haplotypes to clusters in the north of Ireland than to the south, reflecting an overall correlation between Scottish/north English contribution and ChromoPainter PC1 position in Fig 1 (Linear regression: p < 2×10-16, r2 = 0.24).

North to south variation in Ireland and Britain are therefore not independent, reflecting major gene flow between the north of Ireland and Scotland (Fig 5) which resonates with three layers of historical contacts. First, the presence of individuals with strong Irish affinity among the third generation PoBI Scottish sample can be plausibly attributed to major economic migration from Ireland in the 19th and 20th centuries [6]. Second, the large proportion of Northern Irish who retain genomes indistinguishable from those sampled in Scotland accords with the major settlements (including the Ulster Plantation) of mainly Scottish farmers following the 16th Century Elizabethan conquest of Ireland which led to these forming the majority of the Ulster population. Third, the suspected Irish colonisation of Scotland through the Dál Riata maritime kingdom, which expanded across Ulster and the west coast of Scotland in the 6th and 7th centuries, linked to the introduction and spread of Gaelic languages [3]. Such a migratory event could work to homogenise older layers of Scottish population structure, in a similar manner as noted on the east coasts of Britain and Ireland. Earlier communications and movements across the Irish Sea are also likely, which at its narrowest point separates Ireland from Scotland by approximately 20 km.

ireland-britain-genetic-geography
Genes mirror geography in the British Isles. (A) fineSTRUCTURE clustering dendrogram for combined Irish and British data. Data principally split into Irish and British groups before subdividing into a total of 50 distinct clusters, which are combined into cluster groups for clusters that formed clades in the dendrogram, overlapped in principal component space (B) and were sampled from regions that are geographically contiguous. Names and labels follow the geographical provenance for the majority of data within the cluster group. Details for each cluster in the dendrogram are provided in S2 Fig. (B) Principal component analysis (PCA) of haplotypic similarity based on the ChromoPainter coancestry matrix, coloured by cluster group with their median locations labelled. We have chosen to present PC1 versus PC4 here as these components capture new information regarding correlation between haplotypic variation across Britain and Ireland and geography, while PC2 and PC3 (Fig 4) capture previously reported splitting for Orkney and Wales from Britain [7]. A map of Ireland and Britain is shown for comparison, coloured by sampling regions for cluster groups, the boundaries of which are defined by the Nomenclature of Territorial Units for Statistics (NUTS 2010), with some regions combined. Sampling regions are coloured by the cluster group with the majority presence in the sampling region; some sampling regions have significant minority cluster group representations as well, for example the Northern Ireland sampling region (UKN0; NUTS 2010) is majorly explained by the NICS cluster group but also has significant representation from the NLU cluster group. The PCA plot has been rotated clockwise by 5 degrees to highlight its similarity with the geographical map of the Ireland and Britain. NI, Northern Ireland; PC, principal component. Cluster groups that share names with groups from Fig 1 (NLU; SMN; CLN; CNN) have an average of 80% of their samples shared with the initial cluster groups. © EuroGeographics for the map and administrative boundaries, note some boundaries have been subsumed or modified to better reflect sampling regions.

Genomic footprints of migration into Ireland

Quite interesting is that it is haplogroups, and not admixture, that which defines the oldest migration layers into Ireland. Without evidence of paternal Y-DNA lineages we would probably not be able to ascertain the oldest migrations and languages broght by migrants, including Celtic languages:

Of all the European populations considered, ancestral influence in Irish genomes was best represented by modern Scandinavians and northern Europeans, with a significant single-date one-source admixture event overlapping the historical period of the Norse-Viking settlements in Ireland (p < 0.01; fit quality FQB > 0.985; Fig 6). (…) This suggests a contribution of historical Viking settlement to the contemporary Irish genome and contrasts with previous estimates of Viking ancestry in Ireland based on Y chromosome haplotypes, which have been very low [25]. The modern-day paucity of Norse-Viking Y chromosome haplotypes may be a consequence of drift with the small patrilineal effective population size, or could have social origins with Norse males having less influence after their military defeat and demise as an identifiable community in the 11th century, with persistence of the autosomal signal through recombination.

European admixture date estimates in northwest Ulster did not overlap the Viking age but did include the Norman period and the Plantations

The genetic legacies of the populations of Ireland and Britain are therefore extensively intertwined and, unlike admixture from northern Europe, too complex to model with GLOBETROTTER.

ireland-admixture-estimates
All-Ireland GLOBETROTTER admixture date estimates for European and British surrogate admixing populations. A summary of the date estimates and 95% confidence intervals for inferred admixture events into Ireland from European and British admixing sources is shown in (A), with ancestry proportion estimates for each historical source population for the two events and example coancestry curves shown in (B). In the coancestry curves Relative joint probability estimates the pairwise probability that two haplotype chunks separated by a given genetic distance come from the two modeled source populations respectively (ie FRA(8) and NOR-SG); if a single admixture event occurred, these curves are expected to decay exponentially at a rate corresponding to the number of generations since the event. The green fitted line describes this GLOBETROTTER fitted exponential decay for the coancestry curve. If the sources come from the same ancestral group the slope of this curve will be negative (as with FRA(8) vs FRA(8)), while a positive slope indicates that sources come from different admixing groups (as with FRA(8) vs NOR-SG). The adjacent bar plot shows the inferred genetic composition of the historical admixing sources modelled as a mixture of the sampled modern populations. A European admixture event was estimated by GLOBETROTTER corresponding to the historical record of the Viking age, with major contributions from sources similar to modern Scandinavians and northern Europeans and minor contributions from southern European-like sources. For admixture date estimates from British-like sources the influence of the Norman settlement and the Plantations could not be disentangled, with the point estimate date for admixture falling between these two eras and GLOBETROTTER unable to adequately resolve source and proportion details of admixture event (fit quality FQB< 0.985). The relative noise of the coancestry curves reflects the uncertainty of the British event. Cluster labels (for the European clustering dendrogram, see S4 Fig; for the PoBI clustering dendrogram, see S3 Fig): FRA(8), France cluster 8; NOR-SG, Norway, with significant minor representations from Sweden and Germany; SE_ENG, southeast England; N_SCOT(4) northern Scotland cluster 4.

Another study that strengthens the need to ascertain haplogroup-admixture differences between Yamna/Bell Beaker and Sredni Stog/Corded Ware.

Text and images from preprint article under a CC-BY-NC-ND 4.0 International license.

Featured image, from the article on Science Reports: The clustering of individuals with Irish and British ancestry based solely on genetics. Shown are 30 clusters identified by fineStructure from 2,103 Irish and British individuals. The dendrogram (left) shows the tree of clusters inferred by fineStructure and the map (right) shows the geographic origin of 192 Atlas Irish individuals and 1,611 British individuals from the Peoples of the British Isles (PoBI) cohort, labelled according to fineStructure cluster membership. Individuals are placed at the average latitude and longitude of either their great-grandparental (Atlas) or grandparental (PoBI) birthplaces. Great Britain is separated into England, Scotland, and Wales. The island of Ireland is split into the four Provinces; Ulster, Connacht, Leinster, and Munster. The outline of Britain was sourced from Global Administrative Areas (2012). GADM database of Global Administrative Areas, version 2.0. www.gadm.org. The outline of Ireland was sourced from Open Street Map Ireland, Copyright OpenStreetMap Contributors, (https://www.openstreetmap.ie/) – data available under the Open Database Licence. The figure was plotted in the statistical software language R46, version 3.4.1, with various packages.
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