More Celts of hg. R1b, more Afanasievo ancestry, more maps

iron-age-early-celtic-expansion

Interesting recent developments:

Celts and hg. R1b

Gauls

Recent paper (behind paywall) Multi-scale archaeogenetic study of two French Iron Age communities: From internal social- to broad-scale population dynamics, by Fischer et al. J Archaeol Sci (2019).

In it, Fischer and colleagues update their previous data for the Y-DNA of Gauls from the Urville-Nacqueville necropolis, Normandy (ca. 300-100 BC), with 8 samples of hg. R, at least 5 of them R1b. They also report new data from the Gallic cemetery at Gurgy ‘Les Noisats’, Southern Paris Basin (ca. 120-80 BC), with 19 samples of hg. R, at least 13 of them R1b.

In both cases, it is likely that both communities belonged (each) to the same paternal lineages, hence the patrilocal residence rules and patrilineality described for Gallic groups, also supported by the different maternal gene pools.

The interesting data would be whether these individuals were of hg. R1b-L21, hence mainly local lineages later replaced or displaced to the west, or – a priori much more likely – of some R1b-U152 and/or R1b-DF27 subclades from Central Europe that became less and less prevalent as Celts expanded into more isolated regions south of the Pyrenees and into the British Isles. Such information is lacking in the paper, probably due to the poor coverage of the samples.

early-iron-age-europe-y-dna
Y-DNA haplogroups in Europe during the Early Iron Age. See full map.

Other Celts

As for early Celts, we already have:

Celtiberians from the Basque Country (one of hg. I2a) and likely Celtic genetic influence in north-east Iberia (all R1b), where Iberian languages spread later, showing that Celts expanded from some place in Central Europe, probably already with the Urnfield culture (ca. 1300 BC on).

Two Hallstatt samples from Bylany, Bohemia (ca. 836-780 BC), by Damgaard et al. Nature (2018), one of them of hg. R1b-U152.

mitterkirchen-grab-hu-i-8-hallstatt
Photo and diagram of burial HÜ-I/8, Mitterkirchen, Oberösterreich, Leskovar 1998.

Another Hallstatt HaC/D1 sample from Mittelkirchen, Austria (ca. 850-650/600), by Kiesslich et al. (2012), with predicted hg. G2a (see Athey’s haplogroup prediction).

One sample of early La Tène culture A from Putzenfeld am Dürrnberg, Hallein, Austria (ca 450–380 BC), by Kiesslich et al. (2012), with predicted hg. R1b (see Athey’s haplogroup prediction).

NOTE. For potential unreliability of haplogroup prediction with Whit Atheys’ haplogroup predictor, see e.g. Zhang et al. (2017).

kelten-dna-putzenfeld-duerrnberg-grab-376
Photo and diagram of Burial 376, Putzenfeld, Dürrnberg bei Hallein, Moser 2007.

Three Britons from Hinxton, South Cambridgeshire (ca. 170 BC – AD 80) from Schiffels et al. (2016), two of them of local hg. R1b-S461.

Indirectly, data of Vikings by Margaryan et al. (2019) from the British Isles and beyond show hg. R1b associated with modern British-like ancestry, also linked to early “Picts”, hence likely associated with Britons even after the Anglo-Saxon settlement. Supporting both (1) my recent prediction of hg. R1b-M167 expanding with Celts and (2) the reason for its presence among modern Scandinavians, is the finding of the first ancient sample of this subclade (VK166) among the Vikings of St John’s College Oxford, associated with the ‘St Brice’s Day Massacre’ (see Margaryan et al. 2019 supplementary materials).

The R1b-M167 sample shows 23.5% British-like ancestry, hence autosomally closer to other local samples (and related to the likely Picts from Orkney) than to some of his deceased partners at the site. Other samples with sizeable British-like ancestry include VK177 (32.6%, hg. R1b-U152), VK173 (33.3%, hg. I2a1b1a), or VK150 (25.6%, hg. I2a1b1a), while typical Germanic subclades like I1 or R1b-U106 – which may be associated with Anglo-Saxons, too – tend to show less.

late-iron-age-europe-y-dna
Y-DNA haplogroups in Europe during the Late Iron Age. See full map.

I remember some commenter asking recently what would happen to the theory of Proto-Indo-European-speaking R1b-rich Yamnaya culture if Celts expanded with hg. R1a, because there were only one hg. R1b and one (possibly) G2a from Hallstatt. As it turns out, they were mostly R1b. However, the increasingly frequent obsession of searching for specific haplogroups and ancestry during the Iron Age and the Middle Ages is weird, even as a desperate attempt, because:

  1. it is evident that the more recent the ancient DNA samples are, the more they are going to resemble modern populations of the same area, so ancient DNA would become essentially useless;
  2. cultures from the early Iron Age onward (and even earlier) were based on increasingly complex sociopolitical systems everywhere, which is reflected in haplogroup and ancestry variability, e.g. among Balts, East Germanic peoples, Slavs (of hg. E1b-V13, I2a-L621), or Tocharians.

In fact, even the finding of hg. R1b among Celts of central and western Europe during the Iron Age is rather unenlightening, because more specific subclades and information on ancestry changes are needed to reach any meaningful conclusion as to migration vs. acculturation waves of expanding Celtic languages, which spread into areas that were mostly Indo-European-speaking since the Bell Beaker expansion.

Afanasevo ancestry in Asia

Wang and colleagues continue to publish interesting analyses, now in the preprint Inland-coastal bifurcation of southern East Asians revealed by Hmong-Mien genomic history, by Xia et al. bioRxiv (2019).

Interesting excerpt (emphasis mine):

Although the Devil’s Cave ancestry is generally the predominant East Asian lineage in North Asia and adjacent areas, there is an intriguing discrepancy between the eastern [Korean, Japanese, Tungusic (except northernmost Oroqen), and Mongolic (except westernmost Kalmyk) speakers] and the western part [West Xiōngnú (~2,150 BP), Tiānshān Hun (~1,500 BP), Turkic-speaking Karakhanid (~1,000 BP) and Tuva, and Kalmyk]. Whereas the East Asian ancestry of populations in the western part has entirely belonged to the Devil’s Cave lineage till now, populations in the eastern part have received the genomic influence from an Amis-related lineage (17.4–52.1%) posterior to the presence of the Devil’s Cave population roughly in the same region (~7,600 BP)12. Analogically, archaeological record has documented the transmission of wet-rice cultivation from coastal China (Shāndōng and/or Liáoníng Peninsula) to Northeast Asia, notably the Korean Peninsula (Mumun pottery period, since ~3,500 BP) and the Japanese archipelago (Yayoi period, since ~2,900 BP)2. Especially for Japanese, the Austronesian-related linguistic influence in Japanese may indicate a potential contact between the Proto-Japonic speakers and population(s) affiliating to the coastal lineage. Thus, our results imply that a southern-East-Asian-related lineage could be arguably associated with the dispersal of wet-rice agriculture in Northeast Asia at least to some extent.

afanasevo-namazga-devils-gate-xiongnu-huns-tianshan-admixture
Spatial and temporal distribution of ancestries in East Asians. Reference populations and corresponding hypothesized ancestral populations: (1) Devil’s Cave (~7,600 BP), the northern East Asian lineage; (2) Amis, the southern East Asian lineage (= AHM + AAA + AAN); (3) Hòabìnhian (~7,900 BP), a lineage related to Andamanese and indigenous hunter-gatherer of MSEA; (4) Kolyma (~9,800 BP), “Ancient Palaeo-Siberians”; (5) Afanasievo (~4,800 BP), steppe ancestry; (6) Namazga (~5,200 BP), the lineage of Chalcolithic Central Asian. Here, we report the best-fitting results of qpAdm based on following criteria: (1) a feasible p-value (&mt; 0.05), (2) feasible proportions of all the ancestral components (mean &mt; 0 and standard error < mean), and (3) with the highest p-value if meeting previous conditions.

In this case, the study doesn’t compare Steppe_MLBA, though, so the findings of Afanasievo ancestry have to be taken with a pinch of salt. They are, however, compared to Namazga, so “Steppe ancestry” is there. Taking into account the limited amount of Yamnaya-like ancestry that could have reached the Tian Shan area with the Srubna-Andronovo horizon in the Iron Age (see here), and the amount of Yamnaya-like ancestry that appears in some of these populations, it seems unlikely that this amount of “Steppe ancestry” would emerge as based only on Steppe_MLBA, hence the most likely contacts of Turkic peoples with populations of both Afanasievo (first) and Corded Ware-derived ancestry (later) to the west of Lake Baikal.

(1) The simplification of ancestral components into A vs. B vs. C… (when many were already mixed), and (2) the simplistic selection of one OR the other in the preferred models (such as those published for Yamnaya or Corded Ware), both common strategies in population genomics pose evident problems when assessing the actual gene flow from some populations into others.

Also, it seems that when the “Steppe”-like contribution is small, both Yamnaya and Corded Ware ancestry will be good fits in admixed populations of Central Asia, due to the presence of peoples of EHG-like (viz. West Siberia HG) and/or CHG-like (viz. Namazga) ancestry in the area. Unless and until these problems are addressed, there is little that can be confidently said about the history of Yamnaya vs. Corded Ware admixture among Asian peoples.

Maps, maps, and more maps

As you have probably noticed if you follow this blog regularly, I have been experimenting with GIS software in the past month or so, trying to map haplogroups and ancestry components (see examples for Vikings, Corded Ware, and Yamnaya). My idea was to show the (pre)historical evolution of ancestry and haplogroups coupled with the atlas of prehistoric migrations, but I have to understand first what I can do with GIS statistical tools.

My latest exercise has been to map modern haplogroup distribution (now added to the main menu above) using data from the latest available reports. While there have been no great surprises – beyond the sometimes awful display of data by some papers – I think it is becoming clearer with each new publication how wrong it was for geneticists to target initially those populations considered “isolated” – hence subject to strong founder effects – to extrapolate language relationships. For example:

  • The mapping of R1b-M269, in particular basal subclades, corresponds nicely with the Indo-European expansions.
  • There is no clear relationship of R1b, not even R1b-DF27 (especially basal subclades), with Basques. There is no apparent relationship between the distribution of R1b-M269 and some mythical non-Indo-European “Old Europeans”, like Etruscans or Caucasian speakers, either.
  • Basal R1a-M417 shows an interesting distribution, as do maps of basal Z282 and Z93 subclades, despite the evident late bottlenecks and acculturation among Slavs.
  • The distribution of hg. N1a-VL29 (and other N1a-L392 subclades) is clearly dissociated from Uralic peoples, and their expansion in the whole Baltic Sea during the Iron Age doesn’t seem to be related to any specific linguistic expansion.
  • haplogroup-n1a-vl29
    Modern distribution of haplogroup N1a-VL29. See full map.
  • Even the most recent association in Post et al. (2019) with hg. N1a-Z1639 – due to the lack of relationship of Uralic with N1a-VL29 – seems like a stretch, seeing how it probably expanded from the Kola Peninsula and the East Urals, and neither the Lovozero Ware nor forest hunter-fishers of the Cis- and Trans-Urals regions were Uralic-speaking cultures.
  • The current prevalence of hg. R1b-M73 supports its likely expansion with Turkic-speaking peoples.
  • The distribution of haplogroup R1b-V88 in Africa doesn’t look like it was a mere founder effect in Chadic peoples – although they certainly underwent a bottleneck under it.
  • The distribution of R1a-M420 (xM198) and hg. R1b-M343 (possibly not fully depicted in the east) seem to be related to expansions close to the Caucasus, supporting once more their location in Eastern Europe / West Siberia during the Mesolithic.
  • The mapping of E1b-V13 and I-M170 (I haven’t yet divided it into subclades) are particularly relevant for the recent eastward expansion of early Slavic peoples.

All in all, modern haplogroup distribution might have been used to ascertain prehistoric language movements even in the 2000s. It was the obsession with (and the wrong assumptions about) the “purity” of certain populations – say, Basques or Finns – what caused many of the interpretation problems and circular reasoning we are still seeing today.

I have also updated maps of Y-chromosome haplogroups reported for ancient samples in Europe and/or West Eurasia for the Early Eneolithic, Early Chalcolithic, Late Chalcolithic, Early Bronze Age, Middle Bronze Age, Late Bronze Age, Early Iron Age, Late Iron Age, Antiquity, and Middle Ages.

Haplogroup inference

I have also tried Yleaf v.2 – which seems like an improvement over the infamous v.1 – to test some samples that hobbyists and/or geneticists have reported differently in the past. I have posted the results in this ancient DNA haplogroup page. It doesn’t mean that the inferences I obtain are the correct ones, but now you have yet another source to compare.

Not many surprises here, either:

  • M15-1 and M012, two Proto-Tocharians from Shirenzigou, are of hg. R1b-PH155, not R1b-M269.
  • I0124, the Samara HG, is of hg. R1b-P297, but uncertain for both R1b-M73 and R1b-M269.
  • I0122, the Khvalynsk chieftain, is of hg. R1b-V1636.
  • I2181, the Smyadovo outlier of poor coverage, is possibly of hg. R, and could be of hg. R1b-M269, but could also be even non-P.
  • I6561 from Alexandria is probably of hg. R1a-M417, likely R1a-Z645, maybe R1a-Z93, but can’t be known beyond that, which is more in line with the TMRCA of R1a subclades and the radiocarbon date of the sample.
  • I2181, the Yamnaya individual (supposedly Pre-R1b-L51) at Lopatino II is R1b-M269, negative for R1b-L51. Nothing beyond that.

You can ask me to try mapping more data or to test the haplogroup of more samples, provided you give me a proper link to the relevant data, they are interesting for the subject of this blog…and I have the time to do it.

Related

A Song of Sheep and Horses, revised edition, now available as printed books

cover-song-sheep-and-horses

As I said 6 months ago, 2019 is a tough year to write a blog, because this was going to be a complex regional election year and therefore a time of political promises, hence tenure offers too. Now the preliminary offers have been made, elections have passed, but the timing has slightly shifted toward 2020. So I may have the time, but not really any benefit of dedicating too much effort to the blog, and a lot of potential benefit of dedicating any time to evaluable scientific work.

On the other hand, I saw some potential benefit for publishing texts with ISBNs, hence the updates to the text and the preparation of these printed copies of the books, just in case. While Spain’s accreditation agency has some hard rules for becoming a tenured professor, especially for medical associates (whose years of professional experience are almost worthless compared to published peer-reviewed papers), it is quite flexible in assessing one’s merits.

However, regional and/or autonomous entities are not, and need an official identifier and preferably printed versions to evaluate publications, such as an ISBN for books. I took thus some time about a month ago to update the texts and supplementary materials, to publish a printed copy of the books with Amazon. The first copies have arrived, and they look good.

series-song-sheep-horses-cover

Corrections and Additions

Titles
I have changed the names and order of the books, as I intended for the first publication – as some of you may have noticed when the linguistic book was referred to as the third volume in some parts. In the first concept I just wanted to emphasize that the linguistic work had priority over the rest. Now the whole series and the linguistic volume don’t share the same name, and I hope this added clarity is for the better, despite the linguistic volume being the third one.

Uralic dialects
I have changed the nomenclature for Uralic dialects, as I said recently. I haven’t really modified anything deeper than that, because – unlike adding new information from population genomics – this would require for me to do a thorough research of the most recent publications of Uralic comparative grammar, and I just can’t begin with that right now.

Anyway, the use of terms like Finno-Ugric or Finno-Samic is as correct now for the reconstructed forms as it was before the change in nomenclature.

west-east-uralic-schema

Mediterranean
The most interesting recent genetic data has come from Iberia and the Mediterranean. Lacking direct data from the Italian Peninsula (and thus from the emergence of the Etruscan and Rhaetian ethnolinguistic community), it is becoming clearer how some quite early waves of Indo-Europeans and non-Indo-Europeans expanded and shrank – at least in West Iberia, West Mediterranean, and France.

Finno-Ugric
Some of the main updates to the text have been made to the sections on Finno-Ugric populations, because some interesting new genetic data (especially Y-DNA) have been published in the past months. This is especially true for Baltic Finns and for Ugric populations.

ananino-culture-new

Balto-Slavic
Consequently, and somehow unsurprisingly, the Balto-Slavic section has been affected by this; e.g. by the identification of Early Slavs likely with central-eastern populations dominated by (at least some subclades of) hg. I2a-L621 and E1b-V13.

Maps
I have updated some cultural borders in the prehistoric maps, and the maps with Y-DNA and mtDNA. I have also added one new version of the Early Bronze age map, to better reflect the most likely location of Indo-European languages in the Early European Bronze Age.

As those in software programming will understand, major changes in the files that are used for maps and graphics come with an increasing risk of additional errors, so I would not be surprised if some major ones would be found (I already spotted three of them). Feel free to communicate these errors in any way you see fit.

bronze-age-early-indo-european
European Early Bronze Age: tentative langage map based on linguistics, archaeology, and genetics.

SNPs
I have selected more conservative SNPs in certain controversial cases.

I have also deleted most SNP-related footnotes and replaced them with the marking of each individual tentative SNP, leaving only those footnotes that give important specific information, because:

  • My way of referencing tentative SNP authors did not make it clear which samples were tentative, if there were more than one.
  • It was probably not necessary to see four names repeated 100 times over.
  • Often I don’t really know if the person I have listed as author of the SNP call is the true author – unless I saw the full SNP data posted directly – or just someone who reposted the results.
  • Sometimes there are more than one author of SNPs for a certain sample, but I might have added just one for all.
ancient-dna-all
More than 6000 ancient DNA samples compiled to date.

For a centralized file to host the names of those responsible for the unofficial/tentative SNPs used in the text – and to correct them if necessary -, readers will be eventually able to use Phylogeographer‘s tool for ancient Y-DNA, for which they use (partly) the same data I compiled, adding Y-Full‘s nomenclature and references. You can see another map tool in ArcGIS.

NOTE. As I say in the text, if the final working map tool does not deliver the names, I will publish another supplementary table to the text, listing all tentative SNPs with their respective author(s).

If you are interested in ancient Y-DNA and you want to help develop comprehensive and precise maps of ancient Y-DNA and mtDNA haplogroups, you can contact Hunter Provyn at Phylogeographer.com. You can also find more about phylogeography projects at Iain McDonald’s website.

Graphics
I have also added more samples to both the “Asian” and the “European” PCAs, and to the ADMIXTURE analyses, too.

I previously used certain samples prepared by amateurs from BAM files (like Botai, Okunevo, or Hittites), and the results were obviously less than satisfactory – hence my criticism of the lack of publication of prepared files by the most famous labs, especially the Copenhagen group.

Fortunately for all of us, most published datasets are free, so we don’t have to reinvent the wheel. I criticized genetic labs for not releasing all data, so now it is time for praise, at least for one of them: thank you to all responsible at the Reich Lab for this great merged dataset, which includes samples from other labs.

NOTE. I would like to make my tiny contribution here, for beginners interested in working with these files, so I will update – whenever I have time – the “How To” sections of this blog for PCAs, PCA3d, and ADMIXTURE.

-iron-age-europe-romans
Detail of the PCA of European Iron Age populations. See full versions.

ADMIXTURE
For unsupervised ADMIXTURE in the maps, a K=5 is selected based on the CV, giving a kind of visual WHG : NWAN : CHG/IN : EHG : ENA, but with Steppe ancestry “in between”. Higher K gave worse CV, which I guess depends on the many ancient and modern samples selected (and on the fact that many samples are repeated from different sources in my files, because I did not have time to filter them all individually).

I found some interesting component shared by Central European populations in K=7 to K=9 (from CEU Bell Beakers to Denmark LN to Hungarian EBA to Iberia BA, in a sort of “CEU BBC ancestry” potentially related to North-West Indo-Europeans), but still, I prefer to go for a theoretically more correct visualization instead of cherry-picking the ‘best-looking’ results.

Since I made fun of the search for “Siberian ancestry” in coloured components in Tambets et al. 2018, I have to be consistent and preferred to avoid doing the same here…

qpAdm
In the first publication (in January) and subsequent minor revisions until March, I trusted analyses and ancestry estimates reported by amateurs in 2018, which I used for the text adding my own interpretations. Most of them have been refuted in papers from 2019, as you probably know if you have followed this blog (see very recent examples here, here, or here), compelling me to delete or change them again, and again, and again. I don’t have experience from previous years, although the current pattern must have been evidently repeated many times over, or else we would be still talking about such previous analyses as being confirmed today…

I wanted to be one step ahead of peer-reviewed publications in the books, but I prefer now to go for something safe in the book series, rather than having one potentially interesting prediction – which may or may not be right – and ten huge mistakes that I would have helped to endlessly redistribute among my readers (online and now in print) based on some cherry-picked pairwise comparisons. This is especially true when predictions of “Steppe“- and/or “Siberian“-related ancestry have been published, which, for some reason, seem to go horribly wrong most of the time.

I am sure whole books can be written about why and how this happened (and how this is going to keep happening), based on psychology and sociology, but the reasons are irrelevant, and that would be a futile effort; like writing books about glottochronology and its intermittent popularity due to misunderstood scientist trends. The most efficient way to deal with this problem is to avoid such information altogether, because – as you can see in the current revised text – they wouldn’t really add anything essential to the content of these books, anyway.

Continue reading

Official site of the book series:
A Song of Sheep and Horses: eurafrasia nostratica, eurasia indouralica

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).

EDIT (23 APRIL): In Hernández et al. (2018), the TMRCA of R1b-M167 is reported as 3372-3718 ybp:

The youngest sub-branch, R1b-M167, dates to approximately 3.5 kya (95% CI= 2.5-5.3 kya), i.e. even after the Bronze Age.

r1b-df27-m167-europe
Contour (surface) maps displaying the frequencies of Y-chromosome haplogroup and its sub-lineages across Europe and the Mediterranean basin. Modified from Hernández et al. (2018).

NOTE. Admittedly, the maps are mainly based on Iberian samples and certain limited sampling elsewhere, so most of the frequencies displayed in other territories are extrapolated. Since the percentage of R1b-M167 in France is estimated to be ca. 3%, and in Bavaria ca. 5%, the distribution in Central Europe is probably much higher, and around the Mediterranean much lower than represented in them.

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

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

iberia-migrations-celts

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

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

Abstract:

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

Interesting excerpts:

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

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

iberian-adna

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

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

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

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

Also important are the precise subclades:

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


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

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

See also