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Inventing the whites, what hath fog wrought?

One of the first posts on this blog relating to archaeogenetics involved an essay by me involving reflections on the fact that a particular Y chromosomal haplogroup, N1c (N3a now), had a peculiar distribution which ranged from Siberia to Finland. The argument, at the time, was whether it was a lineage which moved east to west (as suggested by the decline of microsatellite diversity in that direction), or whether it moved west to east (as was suggested by the frequency, which was highest in parts of Uralic Europe).

Today we know the general outline of the answer. The N1c lineage seems to have moved westward along the forest-tundra fringe, along with Uralic peoples in general. Genome-wide evidence shows minor but significant affinities with Siberian people among many European Uralic groups, including the Finns, and to a lesser extent Estonians. Though the genome-wide fraction is small in Finns, 5% or less, because this minor component is so genetically different from the generic Northern European ancestry of this group, it shifts Finns off the normal dimensions of variation for Europeans (in addition to the fact that many Finns have been subject to bottlenecks). The fraction is higher in the Sami, and lower in the Estonians.

Additionally, ancient DNA suggests that the arrival of this ‘eastern’ Uralic mediated ancestry seems to date to the early Iron Age. The hypothesis that the Finnic languages were primal to Baltic Europe, is on shaky ground which has cracked open. Rather, the circumstantial evidence is that Finnic languages replaced Indo-European dialects.

A new paper from Estonia as some more detail to the general outline, as well as highlighting some aspects of adaptation. The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East:

In this study, we compare the genetic ancestry of individuals from two as yet genetically unstudied cultural traditions in Estonia in the context of available modern and ancient datasets: 15 from the Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and 6 from the Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). We also included 5 Pre-Roman to Roman Iron Age Ingrian (500 BC–450 AD) (IngIA) and 7 Middle Age Estonian (1200–1600 AD) (EstMA) individuals to build a dataset for studying the demographic history of the northern parts of the Eastern Baltic from the earliest layer of Mesolithic to modern times. Our findings are consistent with EstBA receiving gene flow from regions with strong Western hunter-gatherer (WHG) affinities and EstIA from populations related to modern Siberians. The latter inference is in accordance with Y chromosome (chrY) distributions in present day populations of the Eastern Baltic, as well as patterns of autosomal variation in the majority of the westernmost Uralic speakers [1, 2, 3, 4, 5]. This ancestry reached the coasts of the Baltic Sea no later than the mid-first millennium BC; i.e., in the same time window as the diversification of west Uralic (Finnic) languages [6]. Furthermore, phenotypic traits often associated with modern Northern Europeans, like light eyes, hair, and skin, as well as lactose tolerance, can be traced back to the Bronze Age in the Eastern Baltic.

An admixture analysis shows how ancestry changed over time (from before 2000 BC down to the Middle Ages). The blue fraction associated with Mesolithic Western European hunter-gatherers (WHG) is the majority throughout (though remember that this is an artifactual construct; not necessarily some real ancestral source population). The Estonia_CCC is a representation of the Ceramic Comb Culture individuals, who were dominant in the Baltic after the Mesolithic hunter-gatherers, who seem to have affinities to populations in Western Europe. The CCC individuals seem very similar to Eastern European hunter-gatherers (EHG), whose ancestry had affinities with Paleo-Siberians (Ancient North Eurasians, or ANE), as well as Mesolithic Western Europeans.

There was once a model where CCC mediated the arrival of Finnic languages, but there are two issues now with this framework. First, CCC did not bring N1c. Second, they did not have any Neo-Siberian East Asian ancestry. Rather, their Paleo-Siberian ancestry is deeply nested within the broader West Eurasian set of populations (though some seem to detect reciprocal gene flow between ANE and some proto-East Asian groups). Genes don’t tell us what languages people spoke, but though N1c is not exclusive to Finnic peoples, it’s one of the diagnostic components, along with East Asian ancestry.

After the CCC, the East Baltic region was dominated by Corded Ware Culture (CWC). This culture is affinal in some way to the older Yamna culture. Whereas males in the Yamna culture tended to carry R1b, CWC males were R1a. The Y chromosomes in the paper show massive turnover, so that in the period between 2000 BC and 500 BC R1a was the overwhelming male lineage around the East Baltic. N1c only arrives in large numbers in the Iron Age. One thing that is interesting is to note that N1c is at very high frequencies among Indo-European speaking groups like Lithuanians (the Rurikid lineage is N1c, though Rurik was reputedly a Swede).

It seems clear to me that the Finnic-speaking populations were interacting extensively with Indo-European peoples as they expanded west.  The Finnic-speaking regions of the Baltic were once inhabited by CWC, including southern Finland (the Sami probably absorbed relic hunter-gatherer populations in the north, seeing as they have some unique mtDNA lineages). This is still evident in the frequencies of R1a, which are higher in Finland than in most of Western Europe. In Finland and Estonia, language-shift occurred. The same can not be said for the Latvians and Lithuanians, two Baltic-speaking groups who likely integrated Finnic men to such an extent that N1c is now as common as R1a.

In terms of ancestry, the patterns are starting to become so clear that I think we are near the end of the line. I’ve been pondering the genetic ancestry of Finnic peoples for 17 years on this weblog, but I think at this point there are only details to fill in (thanks to the good ancient DNA climate of Norden, and, Tartu’s genetics researchers).

So here is the sketch. The first people to settle the post-Pleistocene Baltic were hunter-gathers with affinities to the west (WHG). These in their turn were marginalized by hunter-gatherers with affinities to the east (EHG). We don’t know what languages they spoke, but I wouldn’t be surprised if the CCC spoke some sort of Uralic language. At some point, computational methods of linguistic substrate analysis will get much better (there are some linguists who claim that toponyms in parts of Northern Europe give clues to pre-Neolithic languages). Then they, in turn, were replaced by the CWC people. This is important because the CWC introduced the full agricultural package into the eastern Baltic region. Unlike the rest of Europe, including Scandinavia, Early European Farmers (EEF), who descended from Anatolians in the largest part, never colonized the European northeast. The was due to biogeography (I wrote a blog post on this issue years before we had ancient DNA because the climatic patterns were so clear, along with the archeology). The agricultural toolkit of the first farmers was not as well adapted to the climate of this region as elsewhere.

Even in “ancient-DNA-blind” admixture analyses from ~2010 that genome bloggers ran,  something was different between Finns and Swedes even aside from the eastern ancestry. Swedes clearly had a “southern” component in much higher fraction than Finns or even Lithuanians. With hindsight, it is now clear that this southern component was the legacy of the EEF, who never settled en masse in the northeast. But, this component, which is light-green in the plot above, begins to show up among post-CWC populations in Estonia during the Bronze Age, and definitely by the Iron Age. This, in concert with an increase in the blue “hunter-gatherer” ancestry, indicates gene flow from the west.

What this illustrates is that during this period of cultural stability there was a large interaction zone in the Baltic, mediated by trade, which resulted in marriages across populations. As implied in this and other papers, for the CWC peoples this seems to involve the assimilation of women from other groups, as it was highly patrilineal. The women from the west brought both more WHG ancestry, and, EEF. The authors detect that the pattern continued into modern Estonians, who have slightly more EEF than Bronze and Iron Age samples. As agriculture and trade flourished around the Baltic, reciprocal gene flow homogenized what were earlier rather distinct population groups. This was a continuous and stable parameter over the past few thousand years, even without any considerations of perturbations such as the German settlement of the region during the medieval period.

But this gets ahead of ourselves. In the centuries after 500 BC, it seems that post-CWC cultures gave way to Finnic ones. The language is the clearest sign, but the genetic signatures are also clear. But even the Sami are mostly of the same Northern European stock as the other peoples of Norden. The Uralic component was culturally critical, and, looking at the Y chromosomes it looks to have been mediated by mobile paternal lineages. I am not one who knows enough about the paleoclimate or archaeology to hazard a guess what was happening here, but one speculation I have is that the phenomenon may have been rather similar to that which occurred in Greenland, where the Norse were simply unable to compete with the Thule people, whose lifestyle was better suited to the median climate in Greenland. The interactions cannot have been all hostile, as one can judge by the fact that both the Lithuanians and Latvians were extensively penetrated by N1c bearing men who must have arrived relatively late. There are nearly as many N1c lineages in Lithuania as R1a.

This has analogs elsewhere. As I have pointed out, R1a is found in almost all regions of South Asia, across all population classes. The main exceptions outside of the trans-Himalayan fringe are the Austro-Asiatic Munda, who carry high frequencies of Southeast Asian Y chromosomal lineages and notably lack steppe ancestry and R1a. Just as men of Finnic provenance assimilated to the Indo-European language among the Latvian and Lithuanian tribes, so men of Indo-Aryan origin were clearly integrated into non-Indo-European (Dravidian-speaking) societies across South Asia.

Mait Metspalu, a giant in repose

Archaeologists, historians, and folklorists have some work to do. But like Thanos, Mait Metspalu, who has spearheaded so much of this genetic work in Europe’s northeast, can now go farming! His job is done.

But wait! There’s another act here.

The bigger, more surprising, though not entirely so, implication of this paper is that the Nordic phenotype was not brought to the north by a new people, but that it developed in situ through the mixing of peoples. The evidence from this, and other, papers is that Northern Europeans in the Bronze-Age were considerably darker in complexion and mien than they are today. That selection between the Bronze Age and the present has resulted in a sweeping up in frequencies of derived alleles which are strongly correlated with lighter skin, along with selection in other traits considered typical of Northern Europeans, such as the ability to digest milk sugar.

I took one of the supplemental tables and turned them into a chart:

Compare this chart to the one at the top. Between the Bronze Age and the Estonian Middle Ages, and therefore the modern period, the genome-wide changes have been subtle. But for lactase persistence and many of the pigmentation loci, there has been a substantial change without substantial gene flow (and, the East Asian Finnic ancestry likely introduced “dark” alleles, as one can see ancestral copies of SLC24A5 in Finns).

Lactase persistence is interesting because cattle culture in Europe precedes this allele by thousands of years. Likely Pre-Indo-European farmers seem to have utilized cheese (which has lower sugar content).

As for the pigmentation alleles, the standard caveats of predicting past populations on training-sets of the present apply here. But, the genetic character of the East Baltic region seems to have been overwhelmingly in place by the Late Bronze Age. On a genome-wide basis they would be subtly different from modern people in the region, but not substantially so. But, on these salient loci, they do seem quite different. Looking at the detailed SNPs, there are ancestral copies of both SLC24A5 and SLC45A2 variants which are extremely rare in the area today into the Iron Age.

This is not limited to just Estonia. We have two huge datasets from Western Europe, one covering the transition of Britain from Neolithic to Bronze Age, and another, thousands of years in Iberia.

First, let’s look at some SNPs from Britain.

Data from Britain, dashed red line is modern values from 1000 Genomes

CA-BA represents “Copper Age/Bronze Age.” This is the period when British genetic variation basically resembles that of the present day, especially in the north and west of the island (where Germanic period migrations had a minimal impact). A particular haplotype of HERC2 is strongly predictive of having blue eyes. That SNP is an excellent tag. This is the locus where the blue eye haplotype was at very high frequency during the Mesolithic. The frequency of blue eyes predicted increases with the arrival of the Beaker people…but it continues to rise into the present day! The Central European Beaker samples resemble the British ones on their pigmentation loci, just as they do genome-wide.

Now let’s look at the Iberian data.

The selection here is not as strong, but still evident on lactase and SLC45A2. Again, the key is to focus on the Bronze Age, when most of the ancestry of modern Iberians came into focus, with the fusion of EEF, Central and Northern Europeans, and residual WHG ancestry.

We now have some serious temporal transects of phenotypic change inferred from SNPs in very local regions across Europe, in Iberia, in Britain, and now in Estonia. These are very disparate regions, at three points in Europe. But they all seem to suggest the same thing: European populations became depigmented in situ after their overall genome-wide ancestry was established.

A new child of the fields, not the ancient forests

So I guess the question is why? I honestly have no idea. I doubt it’s Peter Frost’s old model because the timing is all off (the European steppe-tundra that plays a big role in his hypothesis also was gone by 4,000 years ago). My idea from nearly a decade ago that it had something to do with the arrival of agriculture and disease isn’t crazy, but I’m a lot less certain about the role of Vitamin D now than I was then.

And the issue around lactase persistence indicates that it’s not limited to the pigmentation genes. There’s lots of selection going on. Both the steppe populations and the Neolithic farming societies practiced cattle culture. In fact, Britain in the centuries before the arrival of the Beaker people had depopulated as it shifted away from intensive farming to agro-pastoralism. But, the steppe peoples do seem to have introduced the allele common in much of Eurasia associated with lactase persistence. It is the same allele found in Northern India. But, the frequency was initially quite low in the ancestral populations.

Overall as one chapter, that of relationships between populations closes, another opens. There’s a lot of innings left in the adaptation game.

Update: A reader of this weblog has pointed out that there is strong evidence that Northern European pigmentation profiles were all over the steppe and forest-steppe by the Bronze Age. Some of the data support that. But when I look closely at the steppe societies such as the Srubna, it is not clear at all that that was the case. For example, a derived SNP at SLC45A2 is close to fixed in modern northern Europeans, rs16891982 (~98%). The frequency is lower in Southern Europeans, closer to ~90%. In the Srubna and related groups, it is closer to 75%. In the Bell Beaker samples from Britain and Central Europe, it is closer to 65%. The frequency is lower in European farmers, but I don’t see the math working proportion-wise for Corded Ware Culture type ancestry with a dilution of EEF leading to a drop from ~100% to 65%.

Additionally, the Estonian CWC samples in the Reich data are ancestral, not derived.

Basically, there was a lot of heterogeneity. Even amongst groups that were similar on genome-wide terms.

(also note that the derived allele was already present at ~25% among Neolithic farmers)

Addendum: Since I put some time into this post, I made this premium. Many of you can read this because you get two free a month, or you are getting around the paywall. But comments are closed if you are not a member. Just so you know if you are confused.

Addendum: I made this a public post again. I’ll work on the Paypal option this weekend.

26 thoughts on “Inventing the whites, what hath fog wrought?

  1. Razib: As implied in this and other papers, for the CWC peoples this seems to involve the assimilation of women from other groups, as it was highly patrilineal. The women from the west brought both more WHG ancestry, and, EEF. The authors detect that the pattern continued into modern Estonians, who have slightly more EEF than Bronze and Iron Age samples. As agriculture and trade flourished around the Baltic, reciprocal gene flow homogenized what were earlier rather distinct population groups.

    Add to this, I thought the section of sex bias in autosome vs X chromosome towards EEF females and steppe males dissolving over time was quite interesting:

    Results and Discussion: “We also tested for sex biases by comparing outgroup f3 statistics calculated on autosomal (A) and X chromosomal (X) data. The high X-to-A ratio of European-early-farmer-related ancestry observed in Estonian Corded Ware Culture decreases over time and disappears by the Middle Ages (Figures S2C–S2F; Data S1).”. (Relevant figures:

    I’ve been interested in this since sex bias signals on the X for Bronze Age populations were introduced by Goldberg. When these signals can be found in CA-BA, are these signals constant over time, post-BA, after the initial admixture?

    In this case and probably all future cases, there’s a lot of noise in these statistics on the X, so hard to be totally confident about this. Looks like there are less than 20000 SNPs per population for even the best quality comparisons, however, quite a lots of ancient populations. But assuming correct they show signals are not constant over time post-BA, even though the uniparental y is fairly constant over time.

    Thinking about why this might be, it might be that this is the sign of male biased absorption of other R1a populations who, even if they were themselves formed by sex biased admixture*, still have a higher autosomal cut of EEF and contribute more to post-CWC populations’ autosome more than X (because male biased admixture). (Add to that probably ongoing sweeping founder effects on the Y, and low level direct admixture from populations who may have no/reversed sex bias admixture).

    You also see in the figures that the autosome seems to be biased towards sharing with European HG relative to the X, and seems to remain so despite subsequent events. If there was subsequent sex bias WHG female into post-CWC cultures, it doesn’t actually seem to be present in the autosome:X signal for whatever reason. This may be due to another complex sex bias admixture scenario.

    There are probably similar questions all over Europe, perhaps even in places that are relatively isolated post-BA, like Ireland.

    *Analogously, if a people were formed in Event 1 by European males+Native American females, they’d have European Y chromosomes and a bias of autosome:X on European ancestry. But if they then mixed back in Event 2 with European females again, they might in extreme scenarios totally lose the autosome:X bias while keeping the originally European Y, and even in cases where the sex bias was somewhat slightly less dramatic in Event 2 than Event 1. Extreme reversals may be unlikely, but these dynamics could make the sex bias signal weak enough to be very hard to detect (since even the most extreme sex bias in one generation is only 66:50). It seems like the X signal can get unstuck from Y shift through these sort of complex scenarios? Particularly this could be the case in societies which are patrilineally centered and might tend to treat a male individual similarly based on patriline, without much regard to his overall ancestry (unlike, e.g. Latin America?).

  2. There are a lot of cases of people in difficult times surviving primarily on the base of one goat or a few cows.
    Obviously, to be able to fully digest milk without problems is a big advantage.
    So is vitamin D “production” with poor food in a cloudy environment.

    I think that these big advantages were further strengthened by social and sexual selection in environments where depigmentation was a clear asset and no burden, unlike some other regions of the world.
    Its not like lighter skin preference was invented by Europeans, its widespread on a global scale in many traditional cultures.

    Its just in Europe environmental and social-sexual selection could combine, based on a lot of mutations introduced by disparate, even though related, colonisers.

    The funny thing is all those ancestral groups might have fostered their own skin lightning, when they finally mixed completely with the Indo-European “mixer”, the panmixed population got a lot of “lighters” to choose from at fairly high frequencies already.

    That was not the case in the more isolated Northern China. Additionally East Asian skin-hair is generally different and they didnt rely on milk products as much.

  3. i used to think the IE induced mixing was a big deal. not sure tho. oca2/herc2 was segregating in the near east. slc24a5 was in SHG and in the caucasus. i don’t know where slc45a2 came from, but it was low freq initially.

    i think it’s selection.

    i used to think the sami were darker than finns cuz they are just more asian. but now i’m not so sure. perhaps the selection env. for the sami, who were not agriculturalists, was very different?

  4. Sure, selection is the main driving force, but you need the variation to start from.

    Sami are such a peculiar group, their ways being so different from other Europeans.

    One big thing is they ate a lot of fresh meat. Actually they were the only European population to stay on the Paleolithic diet.

    The differences between Northern Chinese pigmentation to hunter gatherers at the same or higher altitude might be explained the same way, with lifestyle environmental adaptation plus social and sexual selection.
    Just not at the same level and with different preconditions.

  5. couple comments on the pigmentation profile information. I’m pretty much in agreement with this blogpost on all the points covered (see nothing to dispute!) but would like to add a little for some more context and reference on what the predictions in the paper mean:

    1) Pigmentation profiles from this paper are based on newest Hirisplex 2018 array covered in paper –
    That basically follows the Fitzpatrick skin scale – – with translation of “Fitzpatrick Scale I=Very Pale skin colour, II=Pale, combined III & IV = Intermediate, V=Dark, VI=Dark to Black”.

    (This is the same instrument used in Brace 2019 – – where you had some variation in their prediction for WHGs, with Cheddar Man 9100ybp and La Brana 7000ybp, in Dark/Dark-to-Black, and Loschbour 8000ybp as Intermediate, roughly the same as the classification in this paper of Narva/Kunda as 40% Intermediate, 60% “Mixed/Unpredictable Intermediate-Dark + Dark + Black skin” here).

    2) So this is a scale that although perhaps very visually salient and clinically useful in a European and American context, compresses quite a lot of the world variation in raw reflectance / “M index” terms, as per Fig 1 from “An Unexpectedly Complex Architecture for Skin Pigmentation in Africans”

    (Annotated this Fig to give a rough sense of the amount of that world variation that the four “major locii” – SLC24A5, GRM5-TYR, APBA2 (OCA2), and SLC45A2 – were found to have in Beleza 2013 –

    This isn’t to imply that ancient individuals were not what people might classify as brown in skin tone (which is assuming other variants we don’t know about didn’t adjust the outcome, etc!), but just to note that unless I’m wrong these findings are still pretty compatible with changes in skin reflectance and M index in 50-10 kya period of out of Africa (because the emergence of shifts in the Hirisplex categories seem to only cover a small subset of overall M variation in humanity, or even a subset of variation assuming Khoi-San represent OoA better than some African populations with higher M). Although I know no one is necessarily proposing a jump start change from 10kya after no change since OoA…

  6. Interestingly, and as I’m sure people here will remember, the already-mentioned SHG who came together via a similar fusion of more western and eastern populations were similarly predicted as more depigmented than either of their assumed ancestral populations. More unrelated, they also showed that extra small touch of the east with derived EDAR like modern Finnic speakers do with their extra Nganasan-like component.

    Frost’s theory, if I’m interpreting it correctly as a very long-term generalized phenomenon, doesn’t seem to pan out fully (but there must be something to the specific part that is the sexual selection angle? does blond hair have any other advantages?), with the data apparently reinforcing some basic points made since the earliest aDNA studies studying pigmentation, but it’s still interesting that you got a trend towards that phenotype at least twice since the Mesolithic (though the SHG not nearly as much as moderns) in different contexts around that area. From a quick look, one of the Latvia_MN samples is also predicted as blond and blue-eyed and the Latvia_MN population, as the post brings up, is shifted towards EHG compared to the preceding Latvia_HG, without implying that extra mixing was necessarily the reason.

    I suppose maybe you just had to get the chance mixing of two population elements in the northeast (steppe, HG and/or HG-heavy farmers) who carried derived alleles for all the different features with subsequent selection for whatever reasons for the lighter variants, as mentioned.

    Another interesting aspect in northern Europe is the dichotomy/cline between the more brown-haired but also more red-haired northwest and the much blonder but less red-haired northeast, with Scandinavia being between the East Baltic and the Isles in those features.

    The increase in lactase persistence seems very recent, especially in southern Europe, since other than Iberians, ancient Romans and Greeks also seem totally lactose intolerant apparently. I still have some questions about that since plenty of populations consume decent amounts of non-low-lactose cow’s milk in modern times without major side effects despite being lactose intolerant. It might simply be due to personal ignorance but I can’t understand if selection for it is that important or what caused its sudden, late rise. Even the Yamnayans had pretty low lactose tolerance, from what I recall…

    Some more BA forest steppe data would also be welcome!

    “Update: A reader of this weblog has pointed out that there is strong evidence that Northern European pigmentation profiles were all over the steppe and forest-steppe by the Bronze Age. Some of the data support that. But when I look closely at the steppe societies such as the Srubna, it is not clear at all that that was the case. For example, a derived SNP at SLC45A2 is close to fixed in modern northern Europeans, rs16891982 (~98%). The frequency is lower in Southern Europeans, closer to ~90%. In the Srubna and related groups, it is closer to 75%. ”

    Also, by that time, western ancestry had increased on the steppe and forest steppe even further (the populations weren’t exactly like Yamnaya anymore, but similar to CWC and might have links with it) so maybe we have a similar phenomenon of west meets east?

  7. More unrelated, they also showed that extra small touch of the east with derived EDAR like modern Finnic speakers do with their extra Nganasan-like component.

    it’s literally the same haplotype! (please note that most researchers tho don’t think SHG contributed much at all to modern nordic people)

    Another interesting aspect in northern Europe is the dichotomy/cline between the more brown-haired but also more red-haired northwest and the much blonder but less red-haired northeast, with Scandinavia being between the East Baltic and the Isles in those features.

    this is an old observation. the western pops definitely have more EEF

  8. Razib, yeah, that’s why I thought it was interesting that this kind of move towards the “contemporary” phenotype happened apparently twice/independently with a decent gap between the two.

    There’s definitely a subtle cline in far northern Europe but I wonder specifically why red hair, not just brown hair, ended up being more common on the northwestern end. What do you think?

  9. @egg: It seems like that based on at least SLC24A5, SLC45A2,IRF4, for SHGs

    Though not for Ukraine HGs (similar place to SHGs on WHG->EHG cline on having an intermediate position, but all Ukraine a bit more EHG than SHG have and bit more variability within Ukraine HGs between Mesolithic->”Neolithic” pottery using forager tradition I think).

    SLC24A5 seems to have a good cline somewhat related to EHG ancestry while other variants related to pigmentation are less so (e.g. Ukraine HG groups close to EHG on cline have no really no greater frequency of derived SLC45A2 than Iron Gates group close to WHG, and the SHG and EHG sampling bucked the trend).

    Re; Latvia_MN samples, out of the couple that they test, both are classified “Intermediate” in skin tone (in contrast to earlier HG) and MN1 is the only classified blondie. Now MN1 actually clusters with other Latvia HG in their paper ( and clusters with other earlier Latvian HG in the Eurogenes Global 25 – (IDs: LatviaMN1 – ZVEJ26 = I4627. LatviaMN2 – ZVEJ31 is present under another ID number or not there? I couldn’t find anything online to match up his/her ID under the format David uses in his PCA data to the ones Saag use in their paper). LatviaMN2 is the EHG like sample.

    “Latvia_MN” are diverse with I4627 and I4437 being like Latvia HG, I4435 and I4554 (Combed Ware?) being EHG like and I4436 being intermediate (but closest to EHG).

    Not a bad idea by any means to look at the difference between the EHG ancestry Combed Ware Latvian samples and others, but in this instance the MN sample classified as blonde doesn’t actually seem to be the EHG-like one (rather the dark hair and brown eyes classified one is the EHG-like). Tentatively that seems in keeping with tendencies towards higher frequency of derived SLC45A2 in EHG and lower derived OCA2 in EHG though?

    Like to see what Hirisplex array does with some more of these populations. Not that it is necessarily more accurate.

  10. Matt, thanks for taking a more individual look at these again. Looking at the data again, I think I must have basically momentarily forgotten, for some reason, that the Latvia_HG population and about half of the “Latvia_MN” individuals were already intermediate on that cline (like the predicted blond and blue-eyed MN1 I had in mind) and rather a couple of Latvia_MN individuals were EHG-like instead so your reminder and correction were very welcome.

    If anything that’d make more sense under a supposition that it might have taken some sort of initial mixing of more western and more eastern populations for that kind of phenotype to emerge. Doh.

  11. Basically I had mentally moved these populations all a bit further to the genetic west than they are for some reason so the varied MN group having some more eastern individuals = those are the intermediate ones…

  12. @Egg, ah yes, I tend to mentally place Motala HG as further east and Latvia HG as closer to WHG than they are also (probably an artefact of how Motala have had priority in time as *the* intermediate population). Although also means the Latvia HG who I think of as WHG like and predicted here to generally have darker skin tones also closer to intermediate SHG than I had thought. I suppose there could be some way that bringing together WHG like and EHG like populations could allow for selection towards lighter hair / skin tone (segregating variants between populations), although I wouldn’t bet on it holding up as a pattern.

  13. So the entry of the Finnic peoples into the East Baltic actually resulted in higher rates of light pigmentation. I can clearly remember a decade ago anthrobloggers not unlike this one being convinced that the proto-Finnics were heavily Siberian in ancestry and phenotype. I guess thats what people were hoping to see, to vindicate their old theories based on cephalic index and the likes.

  14. finnish ancestry is pretty restricted. at least distinct to finnics. the population in europe with the highest siberian sami is darkest in that region. the rest europe got light without finnic gene flow. but believe what u want to!

  15. In qpAdm the non-local ancestry in oldest Estonian IA N1c-carrier sample, who wasn’t born in Estonia either, looks like the Sintashta outliers featured in the Narasimhan preprint. Samis have high siberian ancestry from Bolshoy Oleni Ostrov or something similar. Linguistically that was unlikely to have much to do with Estonian Iron Age, and better fits the “Paleo-Laplandic/Lakelandic” mentioned here:

    As for the pigmentation, the Sintashta outliers were probably dark-ish though (but maybe selected for blondness later). Did anyone check the loci?

  16. You misunderstood my point completely. I never claimed that blondism in Europe is inherited from Finnics. I only stated the obvious, the arrival of the proto-Finnics in the East Baltic resulted in lighter pigmentation rather than a more “Siberian” phenotype.
    It was a long due reply to the people who were convinced that the proto-Finnics were heavily Siberian. As someone who has described himself as interested in the ancestry of Finns, I’m sure you’ve come across those theories too.

    I also dont quite get how the Sami are relevant here, as they are not Finnic. Obviously the proto-Uralics were not light pigmented and were Siberian in phenotype. I was specifically talking about the Finnics.

  17. As someone who lives in an area that has been somewhat Saami historically, note that many people who identify as Saami do so on the basis of a Saami grandparent or great-grandparent and local pride. People who are closer to 100 % Saami genetically do indeed look Asian to me, and in more than the pigmentation.

    There is also a belief that the Saami language is not the original language of the Saami people, but a language that came in with eastern people and replaced the original one in historic times. But I am not well informed on that, and might be wrong.

  18. I’m sorry but that sounds a lot like the theory of Sami coming from northern Russia in the 18th century that a lot of Scandinavian supremacists championed for obvious reasons.

    And lets be real here, no Sami looks actually Asian. They can have some Asiatic features, but one would never mistake them for actual Asians or anything but generally what Americans would refer to as “white” or Razibs pet peeve, “caucasian”.

    As a Finn from Lapland I can say that they do have something “off” about them to my eyes, but saying they look Asian is like saying that Afro-Americans look white.

  19. Some plots for another way of looking at how the predicted phenotype changes with date and periodisation category –

    Bronze Age and Neolithic etc are not “bad” categories to use, but I do think since they vary in meaning from different parts of the Old World and contain wide range of dates, I think just plotting raw dates YBP against prediction score is a useful look at things.

    I think these agree that the change in hair and eye phenotype looks dated together from 2000 BCE to 0 BCE, fairly continuous, based on trying to fit a curve.

    The Nganassan % from Chromopainter seems unrelated to hair or eye colour, other than that, as it increases with time as well, tends to be positively correlated in this set with lighter hair and blue eye colour.

  20. Have you looked closer at the Finnish genomes with ancestral variant for SLC24A5? I have yet to come across a Sami sample with one, and the two Finnish samples I’ve looked at both had elevated sharing with South Asians, which in Finland means most likely Romani ancestry. I find it very unlikely that SLC24A5 in Finns is actually preserved and not acquired later.

  21. Took a look at the ALFRED data. Ancestral SLC24A5 freq might be skewed by sample size, the outlying Finnish sample set with 4,5% ancestral variant is small and there’s also a small Orcadian set with 12%(!) ancestral freq.

    The larger n>100 Finnish sets have same ancestral freq as Norwegians (n=400).

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