Copper & Bronze Age Steppe people (PIE) had mixed light and dark pigmentation

[Maciamo]

A very interesting new paper by Wilde et al. 2014 tested three genes (HERC2, SLC45A2 and TYR) associated with skin, eye and hair pigmentation in 63 ancient samples from the Pontic-Caspian Steppe (mostly modern Ukraine) dating from the Chalcolithic and Bronze Age.

The results are relatively surprising. Mutations in SLC24A5, another gene for fair skin related to SLC45A2, has previously been associated with the spread of Neolithic farmers and Bronze Age Steppe people (Proto-Indo-Europeans). This paper tested SLC45A2, a gene that mediates melanin synthesis. The authors found it to be present in only 43% of the samples. Nearly 100% of modern Europeans possess the light skin variant for both the SLC24A5 and SLC45A2 genes.

One gene alone isn't enough to determine skin colour with certainty. But it probably means that the Proto-Indo-Europeans possessed various shades of skin colour. That may be the result of the very composite nature of the Bronze Age Steppe people, who I believe belonged to a mixture of Y-haplogroups indigenous East European R1a, East Anatolian R1b, and other Near Eastern haplogroups (G2a3b1, J2b2, T1a1a) that may have come through the Balkans/Carpathians.

That SLC24A5 mutation was absent from Mesolithic samples from Spain and Luxembourg, although they did possess the HERC2 mutation. What we can say is that fair skin was positively selected in Europeans in the last 5000 years. We still don't know when this mutation reached fixation (nearly 100%) in the European population. It has struck me before that ancient Romans and Etruscans appeared to have had much darker skin than modern southern Europeans. That could mean that fair skin only became a common feature throughout Europe during the Middle Ages, perhaps in part due to the influx of Germanic and Slavic people into the Roman Empire. Fair pigmentation would presumably have been positively selected more quickly in Northern Europe for reasons of adaptation.

The frequency of the two other genes, HERC2 and TYR, came as an even bigger surprise, as a mere 16% and 4% of Bronze Age Steppe people possessed them. Their modern frequency in Europe is, in all fairness (pun intended), also lower than SLC45A2, being found respectively in 71% and 36% of Europeans. Unfortunately I do not have data on their regional distribution in Europe, nor in the Middle East.

The article mentions that the TYR mutation (rs1042602) is highly polymorphic in Europeans. According to the HapMap data, Tuscans (TSI) have a higher frequency of the derived allele than Northwest Europeans (CEU).

The HERC2 mutation (rs12913832) is associated with skin pigmentation, but especially with eye colour (the GG allele corresponding to blue eyes 99% of the time). So blue eyes were present among Proto-Indo-Europeans, but at low frequency. Blue eyes might have been more common among Mesolithic Europeans, at least in the Atlantic and Baltic region. The 16% of HERC2 in the Pontic Steppe would have come from the genes maternally inherited from Mesolithic Europeans (U2, U4, U5).


On a side note, it is highly frustrating that this team of geneticists tested 63 samples from the Yamna and Catacomb and related cultures and did not test any Y-DNA at all ! This could prove once and for all that R1b people spread from the Pontic Steppe and not with Neolithic Near Eastern farmers as so many academic papers have claimed.

Nevertheless, the supplementary information provides a list of all the samples tested with their location and age, and mtDNA haplogroups, which is quite useful. But the study would have been much more interesting if they had also provided the Y-DNA haplogroups for each sample. We would have been able to determine whether there was an ethnic heterogeneity in the Steppe population 5000 years ago. I have always suspected that R1a tribes lived in the northern forest-steppe during the Yaman period, while R1b lived in coastal areas and in the Northwest Caucasus. G2a3b would also be in the Northwest Caucasus, but I am not so sure about J2b2 and T1a1a.

UPDATE:

Of the 47 samples tested for the HERC2 mutation, only four had the GG allele for blue eyes. Two of them were from the Yamna culture (one from Kalinovka I near Samara in the Lower Volga region, and one from Mayaki near Odessa in southwest Ukraine) and two from the subsequent Catacomb culture (both from Novozvanovka II, near Donetsk in eastern Ukraine). Five other samples were AG, three from the Yamna culture and two from the Catacomb culture. All the Chalcolithic samples were AA (brown eyes). The sample size is small, but that could mean that blue eyes were not present in the Pontic Steppe before the Early Bronze Age, and that there was therefore a migration of people from outside the steppes, or a dramatic expansion from one small part of the steppes.

Only three samples had the TYR mutation. There were two from the Yamna culture, including one that was homozygous (AA). The other one was from the Catacomb culture. Once again, none of the Chalcolithic samples were derived.

Interestingly, the AA sample (KAL1) was also from the Kalinovka I site, but that one was AG for the HERC2 gene, not GG like KAL2. Surprisingly that KAL1 individual belong to mtDNA N1a1a (the only one in all the samples), which is more typical of Southwest Asian Neolithic farmers than of PIE speakers. But the SLC45A2 allele (CC) suggests that he/she had dark skin.

Only 22 samples were tested for the SLC45A2 mutation (rs16891982). The European allele for white skin is GG, which was found in 5 Yamna individuals and one Catacomb skeleton. There were seven other samples that were CG, 3 Yamna, 3 Catacomb, and interestingly one Early Chalcolithic too. The Chalcolithic sample was from Vinogradnoe in southern central Ukraine. It was in the same kurgan as a Yamna and a Catacomb sample that were also CG. That could be a sign of unusual continuity in population in that particular area. But it could also mean that the dating is wrong and that it is Yamna too.


Here is the breakdown of mtDNA haplogroups by period. I don't understand why the authors just listed haplogroup U without mentioning at least the top subclade. They had the data. I have added it myself.

Early Chalcolithic

SMY3 : H (rCRS)
SMY4 : H5
SMY9 : H7a1 or H13
SMY11 : T2e
VIN1 : H (rCRS)

Late Chalcolithic

DUR1 : U5a2a
MAJ8 : T2
MAJ9 : W
MOB1 : U5a1
MOB3 : U5a1

Yamna

BEN3 : H33c
KAL1 : N1a1a
KAL2 : H*
MAJ3 : U5a1
MAJ4 : U5b2
MAJ5 : X2h (?)
NIK1 : T1a
NIK7 : H (rCRS)
OLE1 : T2
OLE7 : J2b
OVI2 : K
OVI3 : U/K
PEJ1 : U5a1
PES7 : H1a1 or H5a1j
POD1 : W6
POD2 : T2
POP1 : T2a1b
POP3 : U2e
POP4 : U5a1
RIL3 : K1
SUG2 : I1a
SUG6 : H1, H3 or H6
SUG7 : H (rCRS)
SUG8 : H (rCRS)
TET2 : U4a1
VIN2 : T1a
VIN5 : T1a
VIN12 : T1

Catacomb

KNO4 : U4
LIS1 : U5a1
LIS2 : U4
LIS3 : H2a1
NEV1 : U5a1
NEV3 : H1, H3 or H6
NOZ1 : U4
NOZ2 : U4
PEJ2 : H1 or H13
PEJ3 : H1, H3 or H6
PEJ4 : H1, H3 or H6
PEJ5 : U4
SAC2 : J2b
SUG5 : H6
TEM1 : U4
TEM2 : H (rCRS)
TEM3 : J1b1a1
TEM4 : U5a1
TEM5 : R1a
TEM6 : R1a
TEM7 : U4
TEM8 : U (?)
TET1 : I1d
VIN3 : U5a1
VIN8 : J1b1a1


Points of interest:

- U5a is found in all periods. All the Bronze Age samples are U5a1, which I have described as the main Indo-European subclade of U5 (although of Mesolithic Northeast European origin).

- Haplogroup T is oddly absent from the Catacomb samples. T1a, which I have linked to the spread of PIE people, is found in several Yamna samples. Chalcolithic samples only have T2, including T2e (also found in LBK samples from Neolithic Germany).

- J1b1a1, one of the maternal lineages most strongly associated with the original pre-IE R1b people in the Near East, is found in two Catacomb samples.

- Haplogroups I and W, which are both strongly associated with PIE migrations, are indeed found in Bronze Age samples, and W even in a Late Chalcolithic sample.

- Note the absence of haplogroup V, which has been very elusive in ancient samples from all periods.

- The Early Chalcolithic samples look very much like Near Eastern farmers. I would expect that these lineages (H5, H7, T2e) represent the arrival of G2a3, J2b2 and T into the Pontic Steppe from the Carpathians or Balkans.

- The Late Chalcolithic samples look much more native Europeans, and probably represent the descendants of the Mesolithic inhabitants of the steppes (I2a1 and/or R1a), or new immigrants from the north.

- Apart from one U4a1 sample from Yamna in Moldova, U4 is found exclusively in Catacomb samples and at extremely high frequency (28%). This tends to confirm my hypothesis that Yamna was an R1b-dominant culture or mixed R1a-R1b, and that R1a tribes from the north invaded the Pontic Steppe from central European Russia and the Volga-Ural during the Catacomb culture. U4 strongly correlates with R1a and peaks in the Volga-Ural region today.

- Overall, the Yamna samples look quite similar to those from the Corded Ware culture. Therefore there might have been more R1a in the Yamna culture that I expected. In that case, the bulk of R1b would have been in the Maykop culture or in very coastal parts of the Pontic Steppe. I believe that the J1b1a samples (TEM3 and VIN8) are the most reliable proxy for Y-DNA R1b. The samples are from Vinogradnoe, immediately north of Crimea, and Temrta in southern Russia, between the North Caucasus and Kalmykia (only about 250 km from Maykop). TEM3 was not tested for autosomal DNA, but VIN8 had apparently dark skin. However it is one of the three samples derived for the TYR gene.

 

[Wilhelm]

Since we don't know the autosomal-dna of these samples, we cannot know if they were Asian-admixed, hence the results, since asians don't have these mutations. Im more in favor of population replacement rather than selection, seems more plausible.

 

[Goga]

Once again according to me you're misinterpreting some facts. Who's saying that those Bronze Age Steppe people were Proto-Indo-European at the first place and are not just Indo-Europized natives? If original R1b came from the Eastern Anatolia they could have Indo-Europized the natives of the Steppes. If that was the case you can also count on Y-DNA hg. J2a. Once again you're ignoring this haplogroup in your PIE story. According to me the Maykop folks Indo-Europized the Yamna folks and then all other Pontic-Caspian Steppes natives. It has been proven that the Maykop folks came from Northwest Iranian Plateau. So the ORIGINAL Maykop folks were according to me R1b & J2a. So, original PIE that Indo-Europized peoples of the Steppes belonged mostly to R1b & J2a! J2a was a very imporant haplogroup among the Maykop folks, maybe part of their elite!

 

[Goga]

Since we don't know the autosomal-dna of these samples, we cannot know if they were Asian-admixed, hence the results, since asians don't have these mutations.

Which mutations? I'm West Asian and I'm AA for rs12913832 and CC for rs1042602. So I'm an 'Asian' from the western parts of the Iranian Plateau and I do have these mutations!

edit

and CG for rs16891982

 

[kamani]

I guess these initial steppe people in the Bronze-Age were still nomadic semi hunter-gatherer, hence they had only partially mixed with the European Neolithic farmers, which explains why they were only 43% "white".
Another conclusion is that their number was small and by mixing with a much larger European population, their genes watered down. If we look at the IE languages, the big ones are: Latin, Germanic, and Slavic, all of which could have gotten their current spread fairly "recently". The mysteries are: Albanian, Greek, and Armenian. If we take out "recent" borrowings, Albanian words are a mix of Germanic, Latin, and non-IE words. Greek and Armenian also are partially non-IE. So the IE languages spread in Europe could be more of a post Iron Age event due to the migrations of various Germanic tribes and the rise of the Roman empire.

 

[Aberdeen]

These are fascinating results, but I don't quite know how to interpret them. Is it possible that the samples were taken after the Indo-European migrations began, and that the people sampled were at that time recent incomers to the area? These days, people don't just pack up en mass and emigrate as a complete group, but in a tribal society of the past that could have happened. If these people are representative of the IE folk, I agree with kamani that their genes must have been watered down considerably as their language group and their culture spread through Europe. However, if we look at the timing of when linguists say that different branches of the IE languages hived off, it doesn't really support the idea of it being a recent event, as the most recent major linguistic development (proto-German) apparently developed about 2500 years ago and other IE languages being much older.

 

[Nobody1]

Could it be that the other half of these samples were Mongoloids?
And/Or was SLC24A5 (Rs1426654) also tested?

The Neolithic peoples in Europe (Ötzi / Stuttgart) were already tested positive to have Rs1426654 (A/A) light-skin; In contrast to dark-skinned Mesolithic Hunter-gatherers (Loschbour / LaBrana) Rs1426654 (G/G) which is described [info box on the right] as - 'probably darker-skinned, Asian or African ancestry'
http://snpedia.com/index.php/Rs1426654(A;A)


These Mongoloids/Lappanoids could also explain Brachycephaly in Europe; The non-(pre)-Indo-European Ligurians were examined as proper Brachycephalic (Alpinoid) and identical to the Lapps (Uralic);

Roberto Bosi - The Lapps (1977)
Then Virchow examining a number of Lappish skulls at Helsinki, Lund and Copenhagen, in conjunction with ancient Ligurian skulls, discovered many mutual features suggesting an identical strain....The mandible of the Lapps is always small, the bone/formation unemphasized and the chin of a receding pattern not exactly repeated in any other human group - with the very exception of these almost entirely extinct Ligurians.

So also other pre-(non)-Indo-Europeans -

George Bradshaw - Bradshaw's illustrated hand-book to Switzerland and the Tyrol (1899)
Swiss Lake-dwellings - In his careful investigations of pile dwellings, Dr. Studer met with two extreme types of skulls, the brachycephalic and the dolikoccphalic; the former, at Schaffis and Lüschery (Lake of Bienne), belonging to the pure Stone period, and the latter, at Vinolz and Sutz, to the Bronze period. The facts point to an invasion by the Bronze men, involving a complete transformation of the group of domestic animals; the horse appears for the first time, and new races of sheep and dogs replace the older forms of the Stone period. The occurrence of mesocephalic, and even considerably shortened skulls, in the Bronze period, shows that there was no extinction of the brachycephalic race, but that the two races mixed.

 

[Wilhelm]

Which mutations? I'm West Asian and I'm AA for rs12913832 and CC for rs1042602. So I'm an 'Asian' from the western parts of the Iranian Plateau and I do have these mutations!

edit

and CG for rs16891982

I meant East-Eurasian admixed, part mongoloids that is.

 

[Maciamo]

These are fascinating results, but I don't quite know how to interpret them. Is it possible that the samples were taken after the Indo-European migrations began, and that the people sampled were at that time recent incomers to the area?

The Yamna culture was the first Bronze Age culture in the Pontic Steppe (contemporary with Maykop) and is seen as the source of the PIE migrations.

I see the Catacomb culture as a southern expansion of the Abashevo culture (with a lot of R1a-Z93 lineages) from central European Russia, after the biggest part of the Yamna people left the steppes.

 

[ElHorsto]

I don't see any reason to assume significant mongoloid influence as some suggest. Yamna people probably were mostly west_asian autosomally with slowly increasing north_european over time. The paucity of blue eyes in early IE cultures is not very surprising.

Overall, the Yamna samples look quite similar to those from the Corded Ware culture. Therefore there might have been more R1a in the Yamna culture that I expected. In that case, the bulk of R1b would have been in the Maykop culture or in very coastal parts of the Pontic Steppe.

I would not dismiss the possibility that even R1a is partially of more southern origin like R1b.

 

[Wilhelm]

I don't see any reason to assume significant mongoloid influence as some suggest. Yamna people probably were mostly west_asian autosomally with slowly increasing north_european over time. The paucity of blue eyes in early IE cultures is not very surprising.

Does anyone have the West-ASian or middle-eastern statistics for those pigmentation genes ? Because they onle compare Africa-Europe-Asians...so we can only guess mongoloid admixture,

 

[Sile]

In regards to eye colour, these papers have stated
Sturm et al. (2008) . “A single SNP in an evolutionary conserved region within intron 86 of the HERC2 gene determines human blue-brown eye color.” Am J Hum Genet 82(2):424-31.
Eiberg et al. (2008) . “Blue eye color in humans may be caused by a perfectly associated founder mutation in a regulatory element located within the HERC2 gene inhibiting OCA2 expression.” Hum Genet 123(2):177-87.
Kayser et al. (2008) . “Three genome-wide association studies and a linkage analysis identify HERC2 as a human iris color gene.” Am J Hum Genet 82(2):411-23.

Two copies of the G version of this SNP usually results in blue eyes—but green eyes are possible too, also due to other SNPs. Three studies have found that blue eyes most often result if a person has two copies of the G version of this OCA2-controlling SNP, suggesting that the genetic origin of the trait can be traced back to a single point in human evolution. The authors of one of the studies hypothesize that the mutation event that created this version of the SNP happened somewhere around the Black Sea between about 6,000 and 10,000 years ago. During that time generations of migrating farmers gradually carried agriculture from the Near East to northern Europe, where blue eyes are most often found. These same researchers found that blue-eyed individuals from the Mediterranean (five from Turkey and two from Jordan) also have the G version of this SNP, suggesting that blue eyes the world over can be explained by variation in this one SNP.

The heritability of eye color is extremely high. Twin studies have estimated that 90% to 99% of the variation in human eye color is due to genetics.
In Europeans, 72% chance of blue eyes; 27% chance of green eyes; 1% chance of brown eyes.
Blue eyes are found only in people with European ancestry. The widespread nature of blue eyes in these populations suggests that it was a trait that was somehow selected for after it arose. But there is no known advantage to having blue eyes. In fact, some eye diseases, such as uveal melanoma (the most common eye cancer in adults) and Age-related Macular Degeneration, are more common in people with lighter eyes. The lower rate of such diseases in people with darker eye colors may be due to the protective effects of eumelanin in darker eyes. Just as melanin helps protect skin against UV damage from the sun, melanin may also protect cells in the back of the eye that are important for vision.

You have the GG marker ..........you as in myself

Now the issue is that with all the data above, and the knowledge that G2a and I2a ( apart form the R1 group, which seems not older than the late-neolithic ) has many "european only" clusters, is the only Ydna markers in the northern part of the black sea.
The J , T and L, ydna groups have all no clear clusters without middle-east clusters which leads one to suggest they only arrived in Europe in the early bronze-age.

To conclude with the R1 group, the G2a and I2a seem the only candidates for the above and no J1, J2, L or T migrations to that area until later on

http://dienekes.blogspot.com/2008/05/ancient-y-chromosome-studies.html

 

[Aberdeen]

I want to slap those researchers for not testing the Y DNA. However, I like what the Eurogenes blog had to say about this study.

"Surprisingly, the article doesn't mention Keyser et al. 2009, a very important study which showed that a sample of Kurgan nomads from Bronze and Iron Age South Siberia had frequencies of light hair and eyes comparable to those of present-day Northern and Eastern Europeans. Also worth noting is that the most common Y-chromosome haplogroup among these individuals was R1a, which is today the most frequent haplogroup in Eastern Europe, including Ukraine. What this suggests to me is that the Kurgan cultural horizon was not genetically homogeneous. I suspect that Kurgan groups closer to the Balkans carried significantly higher levels of Near Eastern Neolithic farmer ancestry, and were thus much darker than those in the more temperate northerly regions. However, it seems that at some point, the Neolithic farmer DNA was diluted enough by continuous movements of light pigmented groups from the north and east, possibly made up mostly of males, that there was a major shift in pigmentation traits from Near Eastern-like to Northern European-like across most of Eastern Europe. This scenario actually fits very nicely with the latest on the genetic origins of Europeans."

Some references are provided for further data on the issue. Perhaps these folk were IE, but only one component of the IE folk, with the more northerly component of IE on the steppes being more inclined toward the light complexions and light eye colours that are now common in Europe.

 

[Maciamo]

The only mtDNA from the Bronze Age Pontic Steppe tested before this study were three samples reported in Jeremy Newton's master thesis. These were two C4a6 and one C4a3, all from Odessa region in Ukraine, dating from c. 2000 BCE (Catacomb culture).

There were also seven Neolithic samples from central Ukraine (Dnieper region), among which once again were three C samples (including C4a2), alongside H, U3 and U5a1a.

It's odd that none of the 63 individuals tested by Wilde et al. belonged to haplogroup C when 6 out of 10 of the earlier samples had belonged to C. C4a being mostly a west and central Siberian lineage, I wouldn't be surprised if it was one of the original maternal lineages of Palaeolithic R1 people.

There must have been quite a bit of ethnic heterogeneity in the steppes from the Neolithic to the Bronze Age. That may also explain why the Proto-Indo-Europeans possessed such varied mtDNA haplogroups (see maternal lineages of R1a and maternal lineages of R1b).

 

[Aberdeen]

Here's a link to an abstract of a paper suggesting that the move towards pale skin in Europeans during the last 5000 years is more a result of selection than migration.

www.pnas.org/content/early/2014/03/05/1316513111

I don't know how sound the conclusions are, or whether this paper illuminates the issue or whether it just confuses things. Here's a copy of the abstract.

"Pigmentation is a polygenic trait encompassing some of the most visible phenotypic variation observed in humans. Here we present direct estimates of selection acting on functional alleles in three key genes known to be involved in human pigmentation pathways—HERC2, SLC45A2, and TYR—using allele frequency estimates from Eneolithic, Bronze Age, and modern Eastern European samples and forward simulations. Neutrality was overwhelmingly rejected for all alleles studied, with point estimates of selection ranging from around 2–10% per generation. Our results provide direct evidence that strong selection favoring lighter skin, hair, and eye pigmentation has been operating in European populations over the last 5,000 y."

 

[Alan]

I also believe this is very much a prove of a movement from folks who came from West-South_Central Asian (wider Middle East) into the steppes.

Considering the exclusive high caste distribution of J2a in South Asia and it's general association (after R1 and R2) of it with Indo-Iranic folks, it probably was, together with J2b, part of the Indo European movement too.

@kamani, the mtDNA of the samples look pretty much Neolithic farmer. And considering the fact the Yamna was an animal domestication culture (farming technique), it is unlikely that they were still "pure H&G".

 

[Alan]

I don't see any reason to assume significant mongoloid influence as some suggest. Yamna people probably were mostly west_asian autosomally with slowly increasing north_european over time. The paucity of blue eyes in early IE cultures is not very surprising.



I would not dismiss the possibility that even R1a is partially of more southern origin like R1b.

Exactly, As we know ANE was partly ancestral to Gedrosia-Caucasus/North Euro/Amerindian/ASI. So what if the ANE component among the Indo Europeans was more "Gedrosia_Caucasus like" (note the "like", because Caucasus_Gedrosia as component arose with farmer adfmixture) earlier and drifted through time into a "North European like" component.

 

[kamani]

Here's a link to an abstract of a paper suggesting that the move towards pale skin in Europeans during the last 5000 years is more a result of selection than migration.

www.pnas.org/content/early/2014/03/05/1316513111

good find! it says blue eyes were exclusive to Western Eurasia (fancy word for Europe). This other paper says that blue eyes happened 6000-10000 years ago and are irelevant in terms of evolution; meaning they neither help nor hinder. http://www.sciencedaily.com/releases/2008/01/080130170343.htm. I was fixated that blue eyes are better in ice and snow, because Husky dogs usually have them, but I guess they don't really make a difference (polar bears have brown eyes). Since they didn't spread through evolution, the option left is sexual selection drift (for some reason they looked more attractive to ancient Western Eurasians). I'm not quite sure which one was true thou: "Men like blondes" or "Women like blondes".

 

[Aberdeen]

good find! it says blue eyes were exclusive to Western Eurasia (fancy word for Europe). This other paper says that blue eyes happened 6000-10000 years ago and are irelevant in terms of evolution; meaning they neither help nor hinder. http://www.sciencedaily.com/releases/2008/01/080130170343.htm. I was fixated that blue eyes are better in ice and snow, because Husky dogs usually have them, but I guess they don't really make a difference (polar bears have brown eyes). Since they didn't spread through evolution, the option left is sexual selection drift (for some reason they looked more attractive to ancient Western Eurasians). I'm not quite sure which one was true thou: "Men like blondes" or "Women like blondes".

I think that for reasons of history (many Europeans have blue eyes and Europeans and their descendents overseas have been the dominant force in the world in the last few centuries) there have probably been a lot of people assuming that the Indo-Europeans must have been blue eyed blonds, but in fact the only part of the world where we know that the descendents of the IE folk are still in positions of authority is India. Perhaps the IE folk who swept over Europe were dusky brown eyed people who lost their postions of authority over the centuries because Europe doesn't have a rigid caste system. I'm just not sure what kind of cultural bias would have favoured pale skin, light hair and blue eyes - personally, I like women who are a bit darker and find Arab and Afghan women to be very attractive when not excessively covered up. But perhaps it can be explained by the christian "light is good, dark is bad and evil" bias, which plays into human tendencies to fear the night because we're actually creatures of the day rather than being nocturnal. So perhaps those IE people we're trying to find look different than what we'd expect. Except that we know from looking at the genetics of Hindu brahmins that R1a does appear to have been important in terms of the spread of IE, and most R1a folk in Europe are very stereotypically "white" in terms of skin colour and hair and eye colour. So the more I look at all this, the more confused I feel. Did skin tone and eye colour change while the same haplotypes remained common in eastern Europe?

 

[kamani]

Going along with the idea that original IE people were brown and changed tone once in Europe, the "aryans" of India were never "Nordic white" (they were just an offshot branch of these original IE people). A question is why didn't Selection Drift for blue eyes work in Italy or Spain? Italy has had no significant gene inflow from middle east in the last 2000 years and the paper was saying that the drift could be as fast as 10% per generation. Something doesn't check out..Maybe it wasn't Selection based on looks but based on status; a few blue eyed men became tribal chiefs...This could have only happened later in the Bronze Age and Iron Age, since Neolithic and Early Bronze Age were very egalitarian societies.