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Saturday, May 7, 2022

Population genomics of Stone Age Eurasia (Allentoft et al. 2022 preprint)


Over at bioRxiv at this LINK. It'll take me a few days to read this manuscript properly. Here's the abstract:

The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 11-3 thousand years before present, BP) represent some of the most dramatic lifestyle changes in human evolution. We sequenced 317 genomes of primarily Mesolithic and Neolithic individuals from across Eurasia combined with radiocarbon dates, stable isotope data, and pollen records. Genome imputation and co-analysis with previously published shotgun sequencing data resulted in >1600 complete ancient genome sequences offering fine-grained resolution into the Stone Age populations. We observe that: 1) Hunter-gatherer groups were more genetically diverse than previously known, and deeply divergent between western and eastern Eurasia. 2) We identify hitherto genetically undescribed hunter-gatherers from the Middle Don region that contributed ancestry to the later Yamnaya steppe pastoralists; 3) The genetic impact of the Neolithic transition was highly distinct, east and west of a boundary zone extending from the Black Sea to the Baltic. Large-scale shifts in genetic ancestry occurred to the west of this "Great Divide", including an almost complete replacement of hunter-gatherers in Denmark, while no substantial ancestry shifts took place during the same period to the east. This difference is also reflected in genetic relatedness within the populations, decreasing substantially in the west but not in the east where it remained high until c. 4,000 BP; 4) The second major genetic transformation around 5,000 BP happened at a much faster pace with Steppe-related ancestry reaching most parts of Europe within 1,000-years. Local Neolithic farmers admixed with incoming pastoralists in eastern, western, and southern Europe whereas Scandinavia experienced another near-complete population replacement. Similar dramatic turnover-patterns are evident in western Siberia; 5) Extensive regional differences in the ancestry components involved in these early events remain visible to this day, even within countries. Neolithic farmer ancestry is highest in southern and eastern England while Steppe-related ancestry is highest in the Celtic populations of Scotland, Wales, and Cornwall (this research has been conducted using the UK Biobank resource); 6) Shifts in diet, lifestyle and environment introduced new selection pressures involving at least 21 genomic regions. Most such variants were not universally selected across populations but were only advantageous in particular ancestral backgrounds. Contrary to previous claims, we find that selection on the FADS regions, associated with fatty acid metabolism, began before the Neolithisation of Europe. Similarly, the lactase persistence allele started increasing in frequency before the expansion of Steppe-related groups into Europe and has continued to increase up to the present. Along the genetic cline separating Mesolithic hunter-gatherers from Neolithic farmers, we find significant correlations with trait associations related to skin disorders, diet and lifestyle and mental health status, suggesting marked phenotypic differences between these groups with very different lifestyles. This work provides new insights into major transformations in recent human evolution, elucidating the complex interplay between selection and admixture that shaped patterns of genetic variation in modern populations.

Allentoft et al., Population Genomics of Stone Age Eurasia, bioRxiv, posted May 06, 2022, doi: https://doi.org/10.1101/2022.05.04.490594

See also...

Understanding the Eneolithic steppe

246 comments:

«Oldest   ‹Older   201 – 246 of 246
Simon Stevens said...

@Gaska

“WHGs does not only refer to their geographical location (Western Europe), but also to their autosomal composition. All HGs can be modeled as a mixture of western and eastern HGs, and depending on their percentages they are classified in one or the other category”

That’s completely false, not all Mesolithic/UP Euro HGs can be modeled as a mixture of WHG/EHG. Grotta Continenza, early Mesolithic Sicily HGs (Grotta dell’Uzzo and Grotta d’Oriente), Balma, Chan, Moita, and El Miron (among others) completely lack EHG—some Mesolithic French, Welsh, and Baltic HGs lack it as well. Additionally, EHG and SHG share significantly more alleles with ANE (MA1 and AG3) than WHG. Put simply, EHG affinity can be ascertained if the sample’s genetic profile isn’t adequately captured through other sources, which in the case of late UP/Mesolithic Europe, are the following: Magdalenians, Pinarbasi (AHG), Villabruna (who has minor EHG admixture), Grotta Continenza, and the EM Sicily HGs (Grotta dell’Uzzo/Grotta d’Oriente)—the latter two might be the “purest” WHGs we have.

“it is evident that even in geographically located areas in eastern Europe the HGs were mostly western (Baltic-Lithuania>90 % and Balkans-Iron Gates>85%) and all of them were overwhelmingly R1b (regardless of their subclade).“

The issue is that every time an R1/Q clade appears prominently in a Euro HG group, ANE/EHG autosomal DNA also appear. The oldest P, R, and Q bearers on record, were found in Yana RHS, Mal’ta Buret, and Afontova Gora—ANEs from North Asia. So the UP HGs that occupied Eastern Europe had ANE/R1/Q first (out of all the Euro HGs), because they inhabited the region right next to North Asia—the region where ANEs lived. So did the first R1 and Q bearers travel via airliner, all the way to Iberia, France, and Northern Italy from Siberia 18-28kya—skipping over Eastern Europe in the process? Whether a HG population has more WHG/EHG or less, doesn’t matter. Y-DNA lineages can be prominent in populations with minor/negligible percentages of autosomal DNA from the Y-DNA donor population, just look at Y-DNA N1a in NW Russia/Finland/the Baltic, or R1b-M269 in Assyrians, or R1b-V88 in Central/West SSA tribes. The Finns have 10% or less Siberian admixture, and yet they have close to 60% N1a; Lithuanians have around 30-40% N1a and almost no Siberian admixture. This is called genetic washout. Additionally, you have to specify what Eastern European HGs you’re talking about. Narva and Kunda have varying EHG ranges (24-35% EHG at the Latvian sites vs 0-12% at the Lithuanian sites); other Latvian HGs have EHG levels of 19.7-26.2%, while Neolithic, Eastern Narva HGs are 0-46%. The Baltic_MN/CCC have EHG ranges of 63-100%, while one of them has 19.8% Yamnaya-like DNA; one Latvia_MN outlier has 20.7% EHG, while another has 31.5% Yamnaya-like ancestry. Iron Gates has 15% EHG on average, while the Ukraine HGs were majority Eastern Hunter Gatherer (Ukraine_Neolithic is 49.8%-60.7% EHG vs Ukraine_Mesolithic’s 61.9%-71.4% EHG).

Simon Stevens said...

@Gaska

“The only subclades of R1b older in Ukraine/Russia than in the West are V1636 and Z2103 and neither of them is relevant to the history of L51>L151>P312.”

The oldest P297 bearers on record are from Eastern Europe—they also have EHG autosomal DNA. P297 is the parent clade of both M269 and M73/Y13200, and the oldest R1b-M269 sample is from Chalcolithic Bulgaria (he has around 20% steppe admixture):

Latvia_HG I4630, Zvejnieki, Latvia, 7471-7073 calBCE, mtDNA: U5a2c, Y-DNA: R1b-P297*, MathiesonNature2018

Latvia_HG I4432, Zvejnieki, Latvia, 6072-5920 calBCE, mtDNA: U5a2c*, Y-DNA: R1b-Y13200>Y240021, MathiesonNature2018

Samara_HG I0124, Samara Oblast, Russia, 5660-5535 calBCE, mtDNA: U5a1d, Y-DNA: R1b-Y13200>Y13202, MathiesonNature2015

Bulgaria_C I2181, Smyadovo, Bulgaria, 4606-4447 calBCE, mtDNA: HV15, Y-DNA: R1b-M269, MathiesonNature2018

Additionally, the oldest V2219 and V88 samples are from Eastern Europe:

Serbia_IronGates_Mesolithic I5235, Padina, Serbia, 9221-8548 calBCE, mtDNA: U5b2c*, Y-DNA: R1b-V88>Y127541, MathiesonNature2018

Romania_IronGates_Mesolithic M96, Iron Gates, Schela Cladovei, Romania, 7250-6500 calBCE, mtDNA: U5a1c3*, Y-DNA: R1b-V2219>FTA35720, GonzalezFortesCurrBiol2017

I’d say that P297 and M269 are pretty damn relevant to L51’s history, considering that L51 and Z2103 are descendants of P297>M269>L23. Furthermore, M269, L51, and Z2103 are all associated with the spread of steppe autosomal DNA. P297 and M269 have only been found in Eastern Europe or in EHG admixed samples—M269 is always accompanied with Yamnaya-like ancestry.

“I have already told you many times in order to consider R1b as a typical lineage of the HGs you need samples older than Villabruna, meanwhile Bla bla bla bla bla.”

And I’ve told you this many times:

Villabruna has EHG admixture. He is negative for FGC21012 (V2219 equivalent), P297 equivalents, and V1636 equivalents. He has no calls for L389 either, so he is in no way relevant to M269 or P297. Villabruna dates to 12268-11851 calBCE, while P297 formed from 15600-11800 BCE (median: 13600 BCE), and its TMRCA is 12900-9800 BCE (median: 11300 BCE); L389 formed around 16900-13300 BCE (median: 15100 BCE) and its TMRCA is 15600-11800 BCE (median: 13600 BCE). So if he was relevant, he should have at least carried P297—which existed right before Villabruna—but he didn’t, he didn’t even have calls for L389. R1b-L754 formed around 21300-15500 BCE (median: 18400 BCE), and its TMRCA is 16900-13300 BCE (median: 15100 BCE), so Villabruna lived at least 3300 years after the first L754 bearer. That’s plenty of time for admixture to arrive from ANE admixed, Eastern European Hunter Gatherers (EHGs), you know, the same hunter gatherers who were geographically closest to where the ANEs lived—ANEs are the oldest P/R/Q bearing population on record too. Just because a sample is found somewhere with a particular lineage, doesn’t mean that lineage originated there, especially when we consider archaeology, archaeogenetics, autosomal DNA, formation dates, and the clade’s TMRCA. R1a-YP4132 (formation: 10200 BCE, TMRCA: 4200 BCE) didn’t originate in Kazakhstan with the Andronovo sample I4773 (1463-1568 BCE), despite the fact that he’s the first sample we’ve found with YP4132. So Villabruna man is a genetic dead end with EHG ancestry; he belongs to L754* which is a paragroup. He is not relevant to M269, L754, L389, or P297. Just because he was found in northern Italy with R1b-L754*, doesn’t mean that’s where that line originated. EHGs and SHGs share significantly more alleles with ANEs than WHGs. R1 comes from ANE ultimately, so deductively it makes sense that this clade originated in the most ANE rich groups, located in Eastern Europe, which is the area of Europe closest to where ANEs lived (Siberia/North Asia).

Simon Stevens said...

@Gaska

And here’s more information concerning the EHG ancestry in late Upper Paleolithic/Mesolithic European HGs. The following comes from the paper “Genomic and dietary discontinuities during the Mesolithic and Neolithic in Sicily” (2022):

“Here, we showed that compared to the ~14,000 yBP Epigravettian-associated HG Villabruna in northern Italy, the Sicily EM HGs have a higher genetic affinity to Iberian HGs associated with the Magdalenian and Azilian, such as El Mirón and Balma Guilanyà (Data S2.2). Antonio et al. reported similar affinities for Mesolithic HGs from Continenza (Antonio et al., 2019). Compared to Villabruna, no extra Magdalenian-related ancestry has been found in Continenza or Grotta dell’Uzzo (Data S2.2) (Antonio et al., 2019), while Iberian HGs including El Mirón and Balma Guilanyà have been suggested to carry Epigravettian-related ancestry (Figure S6) (van de Loosdrecht, 2021; Villalba-Mouco et al., 2019). This suggests the Epigravettian-related ancestry in Iberian HGs was likely derived from southern/central Italy. Both archaeological and genomic results suggest a deep connection between the Epigravettian of southern Italy and the Magdalenian and Azilian in Iberia (van de Loosdrecht, 2021), which warrants further investigation”

This suggests that for a “pure” WHG, we should use Continenza or EM Sicily HGs from Grotta dell’Uzzo/d’Oriente, instead of Villabruna, because the latter has EHG.

Simon Stevens said...

Which brings us to this:

“Compared to Sicily EM HGs, Sicily LM HGs derived ~18-21% of their ancestry from an EHG related source, providing evidence for shifting ancestry during the Mesolithic (Figure 3A). After the LGM, foragers associated with Epigravettian assemblages expanded in both Italy and southeastern Europe (Djindjian, 2016; Maier, 2015). Similar to southern Italy, the Balkans were also a glacial refugium. Currently, there are no genomes available for Epigravettian foragers from the Balkans. However, the Mesolithic HGs from this region are among the oldest HGs carrying EHG-related ancestry in southeastern Europe (Figure S6C) and suggest the Balkans as a candidate region for the excess EHG ancestry found in LM Sicily HGs. From the start of the Early Holocene, ∼11,700 yBP onward, an EHG/Anatolia HG (AHG) mixture can be found among HGs from Scandinavia, the Baltic, Ukraine, and the Balkans (Figure S6) (van de Loosdrecht, 2021). This underlines previous reports for a long-standing interaction sphere of HGs with EHG and AHG-related ancestry from northern and eastern Europe toward the Near East and the Caucasus (Feldman et al., 2019; Fu et al., 2016; Mathieson et al., 2018). This population may well have expanded into Sicily in the course of the westward shift of the blade-and-trapeze horizon into central and southern Europe, which seems to have begun after the climatic anomaly around 9,300 yBP (Gronenborn, 2017; van de Loosdrecht, 2021)…In previous studies, the characterisation of the genomic profile of HG typically was based on Villabruna as a proxy for WHG ancestry. However, compared to Villabruna, the Sicily EM HGs showed a higher affinity to many WHG individuals (Data S2.2) and also have a more geographically centered position in a southern refugium. In our model we therefore chose Sicily EM HGs rather than Villabruna as the proxy for WHG ancestry, and found that it helps to differentiate between Magdalenian, EHG and AHG related ancestries in post-LGM and Mesolithic European HGs (Fig. S6, Data S2.6) (van de Loosdrecht, 2021). We also found that, post-LGM Upper Paleolithic and Early Mesolithic Villabruna cluster individuals had diverse genomic ancestries with influences from EHG, Magdalenian-associated HG, or both, rather than forming a genetically homogenous group (Fig. S6). During the Late Glacial, a WHG/EHG mixture was detected for the first time in Bichon (Fig. S6B). This added EHG as an additional dimension to an initial, previously established, Magdalenian-WHG ancestry cline in Iberia and central Europe (Fig. S6A) (Villalba-Mouco et al., 2019). From the start of Holocene, an EHG/AHG ancestry mixture can be found among HGs in northern and (south)eastern Europe (Fig. S6C+D).”

So Villabruna and the Villabruna cluster (including the LM Sicilian HGs) have excess EHG admixture. This explains the presence of R1b-L754* in individuals related to the Villabruna cluster; it also explains the presence of R1b-V88 in younger WHGs.

Simon Stevens said...

@Gaska

And your last actual “point” isn’t a point, it’s a straw man argument. I never said L51 originated in Mongolia, that’s patently absurd. You need to work on your reading/English comprehension. Firstly, the Afanasievo Mongolia male I6222 (3320-2918 calBCE) belongs to a dead end subclade/lineage of R1b-L51 (R1b-L51>P310*/L52*), so him being the ancestor of all L52 males is impossible. Furthermore, I6222 is too young, for L51 formed around 3500-4800 BCE (median: 4100 BCE), and its TMRCA is 3100-4400 BCE (median: 3700 BCE). Secondly, I6222 is almost entirely Yamnaya-like; once again, this proves that wherever there’s subclades of R1b-M269 (L23>L51 and L23>Z2103), there’s steppe/Yamnaya-like/Corded Ware-like autosomal DNA. Put simply, R1b-M269 was spread by the men of the Pontic-Caspian and East European forest steppes. Not one M269 bearer has been found without Yamnaya-like ancestry. I6222 being from Mongolia doesn’t mean shit, for Afanasievo genetically and culturally originated in Eastern Europe. Afanasievo, Corded Ware, and Yamnaya are brother/cousin peoples, hence the shared Y-DNA, mtDNA, and autosomal DNA. All three populations descend from the same hunter gatherer and pastoralist groups of Eastern Europe (especially the ones that inhabited the Pontic Caspian and East European forest steppes).

Copper Axe said...

@Davidski

I've been playing with some of my simulted coordinates and here are some of the results:
https://musaeumscythia.blogspot.com/2022/05/neolithic-don-river-foragers-were-key.html

There seems to be quite some variety in terms of streams of forager ancestry, but I can't say if the percentages themselves are legit, because I don't know how the variety of CHG ancestry in steppe_EMBA samples is reflected on qpadm. If this is largely accurate however, then you could have a scenario where the Yamnaya and Afanasievo populations expanding on the dry steppe recently admixed wtih steppe_en populations, explaining why these groups have a stronger variety than early Corded Ware samples.

I also included a Sredny Stog sample that went from raw file > K13 > G25. I doubt the accuracy on that one due to the conversion process but if it is largely accurate then this sample also perfectly falls within range of the Steppe_EMBA cluster.

Either way, this fully backs up what you have been screaming off the rooftops for a while though.

Matt said...

@Rob and anyone else interested, just one further thing on the tangent about how structured the proto "WHG/Iron Gates HG" pool was...

With the *big* proviso that I'm working with Mesolithic data (which makes generalizing to earlier periods very hazardous), so this is only very loosely relevant to the older data, there are some differences that we can get in f4 stats and qpWave that show that at least by roughly 9000 YBP, HG in Italy and SE Europe don't form a clade to outgroups. Via when comparing the HG dated around 9000 YBP from Gonzales-Fortes 2017 and the HG from Grotta-Continenza at around 10000 YBP from Antonio 2019.

I used Admixtools 2 and the v50 1240k to get precomputed f2 with 572859 SNPs (may have been able to get more by pruning the set more), then compared Romania_IronGates_Mesolithic.SG to Italy_Mesolithic.SG (each of which are sets of 3 HGs who look to have no farmer components in their papers, above), using qpWave and a pRight of Cameroon_SMA, Turkey_Boncuklu_N, Russia_Yana_UP, Sunghir3, Ust_Ishim, Amur_River_Paleolithic (AR33K), Amur_River_LPaleolithic (East Asians), Georgia_Kotias, Iran_Wezmeh, Brazil_Sumidouro.

The results showed that they failed to register as a clade because Italy > related than Romania to Sunghir3, Ust_Ishim, Amur_River_LPaleolithic and Italy < Boncuklu, Kotias, Wezmeh and Brazil_Sumidouro (e.g. Native American with later branching ANE and East Asian). The same pattern was evident in a stronger form testing against Spain_HG (which is Chan and Canes from Gonzales-Fortes; although they're quite different I didn't separate them out in this test, may do later).

Results for both (and some more HG groups): https://imgur.com/a/5BMh8li

Note that the Romania_IronGates_Meso has a different relationship with lots of Near East populations (Boncuklu, and Kotias, and Wezmeh Cave) and also with UP Eurasians (Ust Ishim, Yana and SunghirIII) when compared to the outgroup, so between the two this seems unlikely to be purely due to geneflow SE->Near East? That seems like some clinal pattern (although Romania_Meso, Italy_Meso, Spain_HG marginally fails as a cline in qpWave). (Note also qpWave calculates on all pair stats, not just outgroup, so some stats which are not shown may be driving the relationship; the outgroup stats are just indicative).

Bringing this back to the thread title, there's a similar pattern in the Stone Age Eurasia paper in Extended Fig 5, where they find in their model more UP HG in Italy and its West European descendent pops and more Dzudzuana type Near East HG and ANE in the Romania/Serbia set. (although I believe this is exaggerated by some choices in how they cluster samples).

This is all the Mesolithic though, granted, so we would need more data to see what's happening earlier in time. Very hard to talk about what was the case 15kya or 18kya in Italy vs SE Europe without direct data.

Dospaises said...

@StP "For discussion, I have the L51 phylogenetic tree:

http://www.tropie.tarnow.opoka.org.pl/images/ancient-cwc.l-51malopolska-podkarpacie-czechy.jpg"

What you do mean you have it? Why not just post the YFull link https://www.yfull.com/tree/R-L51/ ?

That is something we all have had access to for a very long time.

Even better is the R-L23 phylogenetic tree at https://www.yfull.com/tree/R-L23/ which shows there are only 3 phylogenetic equivalents in that block and it also shows that both the R-L51 block and the R-Z2103 block are direct descendants of the R-L23 block. Meaning the very first person to have a mutation in the R-L51 block and the very first person to have a mutation in the R-Z2103 block were likely 1st to 3rd cousins since they both descend from people that had all 3 of the mutations in the R-L23 block. Meaning they come from the same clan and same autosomal DNA as well. It would be strange for both of them to have come from a population without Steppe autosomal DNA and all of the sudden all of their descendants carry Steppe autosomal DNA.

capra internetensis said...

@Rob

I don't think Shamanka EBA has enough ANE to work for Okunevo. In G25:
52% Shamanka EBA + 48% Afanasievo - distance 5.83%
43% Shamanka EBA + 32% Tyumen HG + 26% Afanasievo - distance 2.23%

BZK002 (Krasnoyarsk, 2800 BC, Q1b-L330) works nicely:
73% BZK002 + 27% Afanasievo - distance 1.40%

qpAdm models from "Paleolithic to Bronze Age Siberians reveal connections to First Americans and across Eurasia""
2-way (the only one that worked): 76% BZK002 + 24% Steppe EMBA
3-way with highest p (several worked): 36% Baikal LNBA + 47% Botai + 16% Steppe EMBA

Rob said...


Yeah I got that too with G25 set-ups, but maybe I misread something on my qpADM run

Anywahy. here is with BZK002

left pops:
Russia_BA_Okunevo.SG
Russia_Afanasievo
BZK002

best coefficients: 0.311 0.689
SE 0.016
Tail prob 0.38

Right Pops
Cameroon_SMA.DG
Pinarbasi
MA1
WSHG
Israel_PPNB
Brazil_LapaDoSanto_9600BP
Kostenki14a
Tianyuan
Iran_GanjDareh_N
Mongolia_North_N
Russia_HG_Karelia
Satsurblia_HG
Turkmenistan_Gonur_BA_1
Serbia_EN_Starcevo

Rob said...

Anyhow, my point was that is more than the 10-20% estimated in Damgaard, & certainly enough to account for the cultural transmission of Yamnaya-Afansievo traits found in Okunevo

Gaska said...

@SS

Do you realize that you have almost written a book to say nothing new? R1b-L754, R1b-P297, R1b-M73 and R1b-V88 are typical and characteristic lineages of WHGs regardless of their geographical location (France, Italy, Denmark, Spain, Germany, Balkans or Baltic). They are older in these regions than in Ukraine, Russia or Central Asia. Moreover, R1b-M269 is (at the moment) older in Bulgaria, in a typical site of the Gumelnita-Karanovo culture (Old Europe) ergo none of these lineages/subclades has its origin in the EHGs. You can go on thinking that L52 has its origin in Mongolia or you can think that R1b has its origin in Siberia, but the only certainty is that after 10 years you have not found M269>L51>L151>P312 neither in the steppes nor in the forest steppe, nor in Siberia. As long as you do not get it, the steppe theory is only wishful thinking and you are a naive person looking for all kinds of arguments to deny what is evident. Regarding the WHGs, as we have R1b-L754 in Villabruna and as we know that the Villabruna cluster expanded to France (Iboussieres-R1b-L754) and the Iberian Peninsula (El Mirón-18 ka), we will surely find more cases of this lineage in Western Europe, so besides preparing yourself not to suffer a heart attack, prepare new arguments, because saying that P297 is pretty damn relevant to L51's history, and considering that L754 is not, is one of the biggest nonsense I have read in this blog (almost as much as defending that a clearly contaminated sample in Mongolia proves the origin of L51 in the Yamnaya culture, or state that “Whether a HG population has more WHG/EHG or less, doesn’t matter” and then say that “the excess of EHg in Villabruna explains the presence of R1b-L754”).

StP said...

@Dospaises said...
@StP "For discussion, I have the L51 phylogenetic tree:
http://www.tropie.tarnow.opoka.org.pl/images/ancient-cwc.l-51malopolska-podkarpacie-czechy.jpg"
- - What you do mean you have it?
…………
What I mean is that the origins of the L51, L151, and the Czech L151 are discussed here, while in SE Poland we have some concentration of ancient samples from this group, although this is not mentioned.
Linderholm et al. 2020
https://www.nature.com/articles/s41598-020-63138-w.pdf
publishes from SE Poland 19 ancient centralEurCWC, include over 7 R1b CWC! and 2 R1b local BBC! from local CWC R1b .
It probably matters a lot!

Gaska said...

@Wee e said-"Surely the linguistic evidence of Etruscan, not DNA, are what show whether Etruscan was an IE or non-IE language?

Obviously, linguistics is what determines whether a language is Indo-European or not. As far as I know there are only five Non-Indo-European languages written in continental Europe (I have read that Pictish and Ligurian are also Indo-European). The Non-IE are Aquitanian-Basque, Raethian, Etruscan, Iberian and Tartessian. I suppose that NO_IE languages would also exist in the rest of Europe but we do not have written testimonies of them. DNA only serves to check the genetic composition of those peoples who spoke NON_IE languages. In the case of Aquitans and Raethians we still do not have DNA, in the case of Basques, Iberians and Tartessians they are overwhelmingly R1b-P312 and in the case of Etruscans they are mostly R1b-P312 (75%)-Common sense tells us that if we can demonstrate genetic continuity between the BBC (also overwhelmingly P312) and historical peoples like Iberians and Etruscans who we know for sure did not speak IE languages, then it is easy to reason that the BB culture did not spoke an IE language, because there is no reason why those men would change their language in Iberia or Italy and keep it in the rest of Europe. That is, the Iberians and Etruscans can demonstrate genetic continuity at least from the BB culture and yet the supporters of the steppe theory are unable to demonstrate genetic continuity by male line between Yamnaya>CWC>BBC-To this you have to add, that regions where it is known for sure that Indo-European languages were spoken (Greece, Anatolia), there is not a single case of R1b-L51>P312.

Gaska said...

@stp

Thanks for the link

Gaska said...

@Dospaises-

At this point in time when even Max Planck has recognized the inability to prove paternal genetic continuity between Yamnaya>CWC>BBC, do you still think that R1b-L51>L151 has its origin in the Yamnaya culture?

If you are so sure of an eastern origin for this lineage, could you explain us why you believe that this lineage has not yet appeared in those regions?

By the way everyone knows the R1b-L23, L51 Z2103 phylogeny, TMRCA etc.. so you don't need to repeat your posts over and over again.

Rob said...

@ Matt

Right, by 10,000 BC there is Iberian, Italian-WHG, Iron Gates, Dnieper, Baltic, SHG, etc, series of structures.

Ryan said...

Re: admixture between EHG and CHG, I think it's worth noting that most terms marriage relationships are shared between PIE and various Caucasian/Kartvelian languages:

http://loanwords.prehistoricmap.com/wp-content/uploads/2017/06/Bj%C3%B8rn-2017-Foreign-elements-in-the-Proto-Indo-European-vocabulary.pdf

Lots of trade terms too.

Matt said...

https://www.biorxiv.org/content/10.1101/2022.05.15.491973v1.full.pdf -

"Stable population structure in Europe since the Iron Age, despite high mobility" - good batch of from a site in Poland here that is before the conventional date of Slavic expansion. However "The cemetery at Weklice is one of the most important and best studied archaeological sites of Wielbark Culture, which is connected to the Goths and Gepids, on territory of today’s Poland. The cemetery was used from the end of the 1st century to the beginning of the 4th century A.D."

They look a little west of present day Poles in their PCA and overlapping more with Czech possibily, but the PCA may be compressed and doesn't have specific Balto-Slavic vs Western European dimensionality. Will be interesting to see how much Balto-Slavic drift they have when the samples drop (will be under PRJEB52852 at ENA when the paper is published). No y-dna data in the paper unfortunately, which could have been useful to know.

Davidski said...

Yep, waiting for the data from the Polish study.

Impossible to be sure of anything based on the analysis in the preprint.

André de Vasconcelos said...

First time we get our hands on Portuguese samples dated to the Roman Period, including two samples from Conimbriga which seem rather IA-like. This is very exciting even though I suspect we'll pretty much see the same thing we've seen in Spain.

Arza said...

Re: Stable population

BAM files:
https://pastebin.com/m4pHxMuG

They should be available soon.

BTW Genomes from Verteba Cave are online too if anyone is willing to process them (fastq):
https://www.ebi.ac.uk/ena/browser/view/PRJEB38797?show=reads

Ric Hern said...

Got my results at long last. Y-DNA R1b L21 and MtDNA Haplogroup H2a2a1d. Hello to all my Steppe long lost cousins. Heheheeh.

Ric Hern said...

I see a sample from Çatalhöyük 8590 ybp. had MtDNA Haplogroup H2a2a1d and some Romans in Italy...but that was all I could find.

ambron said...

Arza, nice to see you again!

Richard Rocca said...

Gaska said "You have changed Steppe-Yamnaya to Forest Steppe-CWC, but you still can't find R1b-M269>L51. I wish you luck, you're going to need it."

Well, I changed in 2018, which is what unbiased people do when new data is presented. And by the way, I already have knowledge that L51 was there. When you see the samples, you will be very, very disappointed.

Richard Rocca said...

Romulus said... "Rocca you are an Etruscan"

I am thrilled that my U152>L2 subclade has been found in an Iron Age Latin and an Iron Age Etruscan. So yes, I won the Y-DNA lottery. I'm sure you meant it at a slight, but then again, you don't seem all that intelligent.

Romulus the I2a L233+ Proto Balto-Slav, layer of Corded Ware Women said...

It boggles my mind how you will let spaghetti retard make these ridiculous statements about winning the genetic lottery (all the genes made him was short and greasy) , and insult me, but not even approve at least my Fredo comment.

Davidski said...

@Romulus

Yeah, right, see that's why I block some of your comments.

AWood said...

@Rob

People move around and die out, especially in nearly inhabitable landmasses. It's like finding a needle in a haystack. You're not going to argue that 24,000 years ago in Siberia we have someone pretty close to the R1 founder, who is our sample ANE population are you? If the 1000 BC mummies are most closely related to this ancestor + other stuff. Obviously the descendants were in Mongolia and Kazakhstan. Who knows, and who really cares their exact trajectory across Asia between 24,000 - 1000 BC. The minor details are actually not that important, because a reasonable person would say they lived in those territories at some point, even if most died out.

Simon_W said...

I found this finding of the paper fascinating:

"Interestingly, loci associated with overdispersed mood-related polygenic phenotypes recorded among the UK Biobank individuals (like increased anxiety, guilty feelings, and irritability) showed an overrepresentation of the Anatolian farmer ancestry component"

From Vonderach's book on the psychology of peoples I gained the impression that higher excitability (alongside passionateness) is the trait most obviously associated with Basal Eurasian admixture, that is, it is more expressed in South Europeans than in the North, and even more in Arabic and North African people than in South Europeans. This was Vonderach's conclusion after reading lots of stuff about the mentalities of peoples. But he also cited more recent, more academic studies about psychological tests on different populations. Contrary to expectations, a higher extroversion wasn't the trait most clearly associated with South Europeans; instead a higher irritability was this trait.

Now the politically correct explanation for such mentality differences between peoples is that it's entirely cultural, it's acquired, it has no basis in the blood, in the DNA. And that it's a random product of the course of history, maybe influenced by the natural environment at most. But now we see that higher irritability is indeed inherited from the Anatolian farmers. Wow!

Simon_W said...

Re: the brachycephalic Borreby skulls, this is another remarkable surprise. Because for a very long time they were believed to be Mesolithic survivors, descendants of local hunter-gatherers. Now it turned out they were post-CWC and closely related with Bell Beakers, what a difference!

Simon_W said...

If I may add an observation on a falsification of physical anthropological typology in this paper: It's the finding that Steppe ancestry in the British Isles is strongest on the Celtic fringe, and particularly in Ireland, while Anatolian Farmer ancestry is strongest in the south and east of England. Anthropological typologists like Deniker, Eickstedt, Biasutti, Lundman, Coon etc. characterized the Celtic fringe as Mediterranean admixed, as Atlanto-Mediterranean, or even Paleo-Atlantid. Kind of suggesting that Neolithic farmers, maybe WHG-admixed, left a stronger imprint there. But genetically that doesn't seem to be the case.

Simon_W said...

I hope they will improve the resolution of their yDNA haplotree graphs in the supplement part 1. They are completely useless the way they are now, the resolution is so low that you can't read anything.

alex said...

Simon_W said...
"Now the politically correct explanation for such mentality differences between peoples is that it's entirely cultural, it's acquired, it has no basis in the blood, in the DNA. And that it's a random product of the course of history, maybe influenced by the natural environment at most. But now we see that higher irritability is indeed inherited from the Anatolian farmers. Wow!"

And what about the North-South difference in depression and suicides? Is that "in the blood" as well?

Simon_W said...

Which countries have the highest suicide rates? I'm not sure, but isn't it something like Finland or Japan? In any case rather not the Mediterranean countries. So this isn't associated with irritability or excitability, but rather with a difficulty of enjoying life, a lack of positive feelings.

Simon_W said...

And I think it's the rather "stoic" temperaments that tend to be susceptible to this.

Synome said...

@Simon_W

I wouldn't read much into the behavioral associated loci. There are too many confounding factors to use associations from a modern UK sample and try to project that millennia into the past and expect it be meaningful. Time and again these effects have been shown to be due to population stratification and other extraneous factors.

Matt said...

@Simon_W; that seems odd to me. In UK Biobank, where these loci are ascertained, steppe ancestry is enriched in the North of England, Scotland, Wales and Northern Ireland, where anxiety disorders also tend to be enriched - https://ocsi.uk/wp-content/uploads/2020/09/MentalHealth_mood_disorders.png

It's also at odds with some of the geography of self repoted trait Neuroticism in the UK - https://www.bbc.co.uk/news/magazine-31816926 (Excitable Celts and stoic English?)

Matt said...

@Simon, also related, check out - https://www.biorxiv.org/content/10.1101/457515v1.full. The polygenic scores for some of these psychiatric problems are higher in the steppe ancestry rich regions of the UK: https://imgur.com/a/q9QHTIl

Simon Stevens said...

@Gaska

You need to learn how to read English properly, because you clearly can’t comprehend anything I wrote. All you do is construct straw mans.

“R1b-L754, R1b-P297, R1b-M73 and R1b-V88 are typical and characteristic lineages of WHGs regardless of their geographical location (France, Italy, Denmark, Spain, Germany, Balkans or Baltic). They are older in these regions than in Ukraine, Russia or Central Asia.“

Citation needed, prove it, provide the samples. Regardless, the oldest P297, M269, Y13200/M73, V2219, and V88 bearers are EHG admixed HGs from Eastern Europe:

Latvia_HG I4630, Zvejnieki, Latvia, 7471-7073 calBCE, mtDNA: U5a2c, Y-DNA: R1b-P297*, MathiesonNature2018

Latvia_HG I4432, Zvejnieki, Latvia, 6072-5920 calBCE, mtDNA: U5a2c*, Y-DNA: R1b-Y13200>Y240021, MathiesonNature2018

Samara_HG I0124, Samara Oblast, Russia, 5660-5535 calBCE, mtDNA: U5a1d, Y-DNA: R1b-Y13200>Y13202, MathiesonNature2015

Bulgaria_C I2181, Smyadovo, Bulgaria, 4606-4447 calBCE, mtDNA: HV15, Y-DNA: R1b-M269, MathiesonNature2018

Serbia_IronGates_Mesolithic I5235, Padina, Serbia, 9221-8548 calBCE, mtDNA: U5b2c*, Y-DNA: R1b-V88>Y127541, MathiesonNature2018

Romania_IronGates_Mesolithic M96, Iron Gates, Schela Cladovei, Romania, 7250-6500 calBCE, mtDNA: U5a1c3*, Y-DNA: R1b-V2219>FTA35720, GonzalezFortesCurrBiol2017

“R1b-M269 is (at the moment) older in Bulgaria, in a typical site of the Gumelnita-Karanovo culture (Old Europe) ergo none of these lineages/subclades has its origin in the EHGs.”

I2181 has EHG and steppe autosomal DNA, so try again; nothing you said refutes the genetic evidence. From MathiesonNature2018: “In two directly dated individuals from southeastern Europe, one (ANI163) from the Varna I cemetery dated to 4711-4550 BCE and one (I2181) from nearby Smyadovo dated to 4550-4450 BCE, we find far earlier evidence of steppe-related ancestry (Figure 1B,D).” When they modeled him in D stats (Mbuti.DG, CHG, Balkans_Chalcolithic, Balkans_Chalcolithic_outlier) they found the following: “Varna_outlier has Steppe ancestry. Balkans_Chalcolithic_outlier may have steppe ancestry but has no evidence of CHG component (however number of SNPs is low).” In the paper’s supplement, I2181 has 46.1% Yamnaya in qpAdm, with a standard deviation of 17.4%. In admixture analysis, he picked up EHG, and a larger Yamnaya_Samara component. I2181 and the Varna outlier are both on the Khvalynsk cline, unlike the other Balkans Chalcolithic samples: https://3.bp.blogspot.com/-gpo-jpYFsY0/WST-FhAWHMI/AAAAAAAAFpU/VISsvervHfQ34fSBLM7c1KOIkLMAxxhTwCLcB/s1600/Steppe_clines.png

Simon Stevens said...

@Gaska
“You can go on thinking that L52 has its origin in Mongolia or you can think that R1b has its origin in Siberia, but the only certainty is that after 10 years you have not found M269>L51>L151>P312 neither in the steppes nor in the forest steppe, nor in Siberia.”

You are so slow and dense. There is this thing in biology and genetics called phylogeny. R1b derives from R1, R1 comes from R, and R comes from P; the oldest P, Q, and R bearers are the ANE samples from Afontova Gora, Mal’ta Buret, and Yana RHS. ANE DNA arrived in Eastern Europe first, because North Asia is right next to Eastern Europe. And hold on, it’s now P312? First it was M269, then L51, and now you’re shifting the goal posts to P312. Even if P312 originated in Central Europe, it still traces back to a Eastern European, steppe derived source, that being M269 and its subclades (PF7562, L23, Z2103, and L51). Show me one M269 bearing sample without steppe ancestry (EHG + CHG), and that means any subclade of M269, which in case you don’t understand, means L51, L52, Z2103, P312, etc.

And I said the following: “the Afanasievo Mongolia male I6222 (3320-2918 calBCE) belongs to a dead end subclade/lineage of R1b-L51 (R1b-L51>P310*/L52*), so him being the ancestor of all L52 males is impossible. Furthermore, I6222 is too young, for L51 formed around 3500-4800 BCE (median: 4100 BCE), and its TMRCA is 3100-4400 BCE (median: 3700 BCE). Secondly, I6222 is almost entirely Yamnaya-like; once again, this proves that wherever there’s subclades of R1b-M269 (L23>L51 and L23>Z2103), there’s steppe/Yamnaya-like/Corded Ware-like autosomal DNA. Put simply, R1b-M269 was spread by the men of the Pontic-Caspian and East European forest steppes. Not one M269 bearer has been found without Yamnaya-like ancestry.”

Nothing I said endorses an origin in Mongolia. Afanasievo, Yamnaya, and Corded Ware share a common ancestry in Eastern Europe. When these groups branched out, they took with them shared lineages, which is why you have Q1b-FT380500, R1b-Z2103, and R1b-L51 in CWC and Afanasievo—R1b-V1636 exists in Corded Ware and Yamnaya too. Moreover, I6222 belongs to a subclade of L51, that being P310. P310 is a SNP at the L52 level, so I6222 Is R1b-L51>P310*/L52*. That’s a paragroup, so L52 bearing men don’t descend from I6222 (he’s a genetic dead end), however, they share an L51 bearing male ancestor with him, one who lived in Eastern Europe—specifically the East European forest and/or Pontic-Caspian steppes—around 3100-4400 BCE (median: 3700 BCE). This is very straight forward and easy to comprehend. Oh and another Afanasievo male belongs to R1b-L52* (sample ID: C3341); so I6222 being L52*/P310* is even more concrete, because we now have two L52* bearers.

“As long as you do not get it, the steppe theory is only wishful thinking and you are a naive person looking for all kinds of arguments to deny what is evident.”

You’ve provided exactly zero counter evidence.

Simon Stevens said...

@Gaska

“Regarding the WHGs, as we have R1b-L754 in Villabruna and as we know that the Villabruna cluster expanded to France (Iboussieres-R1b-L754) and the Iberian Peninsula (El Mirón-18 ka), we will surely find more cases of this lineage in Western Europe, so besides preparing yourself not to suffer a heart attack, prepare new arguments, because saying that P297 is pretty damn relevant to L51's history, and considering that L754 is not, is one of the biggest nonsense I have read in this blog”

That’s the best you could put together dumbass lol? El Miron doesn’t have EHG, Villabruna has EHG autosomal DNA and R1b-L754* (an EHG derived lineage), which is a paragroup, which means Villabruna man is a paternal dead end. He didn’t leave any male descendants with L389, V1636, or P297; he is negative for FGC21012 (V2219 equivalent), P297 equivalents, and V1636 equivalents—he has no calls for L389. This brings us back to phylogeny, a biological concept you seem to have no understanding of. So let’s go over this once again. Villabruna dates to 12268-11851 calBCE, P297 formed around 15600-11800 BCE (median: 13600 BCE), and its TMRCA is 12900-9800 BCE (median: 11300 BCE). L389 formed around 16900-13300 BCE (median: 15100 BCE) and its TMRCA is 15600-11800 BCE (median: 13600 BCE). So if Villabruna was relevant to R1b-L754>L389>P297>M269, he should have at least carried P297—which existed before Villabruna—but he didn’t, he didn’t even have calls for L389. R1b-L754 formed around 21300-15500 BCE (median: 18400 BCE), and its TMRCA is 16900-13300 BCE (median: 15100 BCE), so Villabruna lived at least 3300 years after the first L754 bearer. Just because a sample is found somewhere with a particular lineage, doesn’t mean that lineage originated there. We have to consider the following evidences in tandem: archaeology, archaeogenetics, autosomal DNA, and phylogeny (which includes subclade data, formation dates, and TMRCAs).

And Iboussieres31 has around 7.7% EHG. From Kale: “so I ran Iboussieres31 F3 outgroup stats to 53 phylogenically significant populations. The most salient observations are as follows...Pretty low affinity across the board, especially as relates to other WHG-EHG cline populations. Not sure how much of that is legit or because of poor quality. Depending on the context of the PCA, this may account for the unusual position. Striking resemblance to the slightly less crappy quality 'Germany_Falkenstein_HG'. The usual method I use to calculate ANE (F3: Outgroup Test MA1 - F3: Outgroup Test Yana / F3: Outgroup AG3 MA1 - F3: Outgroup AG3 Yana) gives 7.7%, which is actually on the low side of things for WHG. This is corroborated by low relative affinity to Tarim_EMBA1 and Botai also.” As stated previously, Iboussieres31 is very low/poor coverage (80k SNPs); he could only be discerned to R1, but he had no negative calls for L754, so he likely belongs to the same L754* paragroup/subclade as Villabruna man. So that’s more of your Basque schizo bullshit debunked.

Simon Stevens said...

@Gaska

And we’ve gone over I6222 dozens of times. He has nothing but derived calls at the M269, L51 and L52 levels, reliable calls found in no other haplogroups/clades. Contamination is measured via the mtDNA and the X-chromosome, not the other autosomes and Y-DNA. If he were contaminated he would have had further calls for more recent clades (anachronistic TMRCAs) beyond the L52 level, but surprise, he doesn’t. We would also expect I6222 to have an inconsistent, anachronistic autosomal profile (due to contamination), but alas, he clusters with all the other Afanasievo_Mongolia samples, and that DNA profile doesn’t exist anymore, so this also discounts contamination. And now we have another Afanasievo sample, one who belongs to the exact same R1b-M269>L51>L52*/P310* clade (sample ID: C3341). So sad, time for you to get another hobby, maybe some English lessons?

Neo said...

New HG sample helps explain low-CHG, high-HG of Corded Ware. Can't wait for the deeper analysis to come.

Samuel Andrews said...

@Ric Hern,

Welcome to the R1b L51>P312>L21 tribe! I say this because Indo-European Y DNA really is the marker of specific IE tribes. Western Corded Ware (L51), then Bell Beaker (P312), then British Bell beaker (L21), etc. Our Y DNA isn't from a line of random men, it comes from a line of whole tribes. Pretty cool.

I can take a look at your mtDNA. I am an mtDNA expert. I especially suggest this because often carries of mHG H are falsely labelled as H2a2a1. You can email me your mtDNA raw data at thepopulationgeneticschannel@gmail.com.





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