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Monday, January 27, 2014

A Mesolithic genome from Spain

Nature today published a paper on the complete genome of La Brana 1, a Mesolithic hunter-gatherer from Iberia: Olalde et al. 2014. Based on genetic variants associated with pigmentation traits, it's likely that this individual had blue eyes, dark hair and deep brown skin.

Moreover, he was probably lactose intolerant (in other words, unlike most Europeans today, he couldn't drink milk as an adult), and his Y-chromosome belonged to the European-specific, but today extremely rare, haplogroup C6 (aka. C-V20), and mtDNA to haplogroup U5b2c1, which again is a European-specific marker. Below is an artist's impression of his mug (courtesy of CSIC), and below that the paper abstract.

Ancient genomic sequences have started to reveal the origin and the demographic impact of farmers from the Neolithic period spreading into Europe1, 2, 3. The adoption of farming, stock breeding and sedentary societies during the Neolithic may have resulted in adaptive changes in genes associated with immunity and diet4. However, the limited data available from earlier hunter-gatherers preclude an understanding of the selective processes associated with this crucial transition to agriculture in recent human evolution. Here we sequence an approximately 7,000-year-old Mesolithic skeleton discovered at the La Braña-Arintero site in León, Spain, to retrieve a complete pre-agricultural European human genome. Analysis of this genome in the context of other ancient samples suggests the existence of a common ancient genomic signature across western and central Eurasia from the Upper Paleolithic to the Mesolithic. The La Braña individual carries ancestral alleles in several skin pigmentation genes, suggesting that the light skin of modern Europeans was not yet ubiquitous in Mesolithic times. Moreover, we provide evidence that a significant number of derived, putatively adaptive variants associated with pathogen resistance in modern Europeans were already present in this hunter-gatherer.

Indeed, the pigmentation traits are basically the same as those of Loschbour, a Mesolithic genome from Luxembourg, featured recently in the groundbreaking Lazaridis et al. preprint (see here). So we can already speculate with some confidence that this was a common, and perhaps dominant, trait combination among European hunter-gatherers.

However, early European farmers, whose ancestors almost certainly migrated to Europe from the Near East during the Neolithic, probably had somewhat different pigmentation traits. We know this because a 7,500 year-old Linearbandkeramik (LBK) farmer genome from Stuttgart, Germany, also featured in Lazaridis et al., showed markers for brown eyes, dark hair, and relatively light skin.

So as things stand, it appears that Europeans only acquired their present coloring, including pale skin and a high incidence of light eyes, relatively recently, well after the hunter-gatherers and farmers began mixing, and their hybrid DNA had time to go through some really powerful selective sweeps. These sweeps were possibly in part a reaction to the Neolithic diet, rich in carbohydrates but poor in vitamin D, amongst other things. Vitamin D doesn't have to be acquired from food because the body can synthesize it from the sun, but this is done more effectively by people with fair skin, giving them an advantage, especially in places like Europe, which has fairly long winters and lots of cloud cover.

But perhaps this isn't the full story, and present-day European pigmentation traits are also sourced from a late migration into Europe of a prevailingly blond people from somewhere in what is now Russia?

This might sound far fetched, but during the middle Bronze Age the Eurasian steppe was home to the Andronovo culture, with archeological links to earlier cultures in what is now southern Russia. Based on the DNA of Andronovo nomads from Kurgans in South Siberia, it seems they had fair skin and a lot of blue eyes and blond hair (see here). They also overwhelmingly belonged to Y-chromosome haplogroup R1a1a, which is very common today in Central and Eastern Europe and also parts of Scandinavia. So it'll be interesting to see the pigmentation markers of Mesolithic Eastern Europeans and Central Asians when their genomes become available, probably in the not too distant future, and if they contributed any ancestry to present-day Europeans. Early indications are that they did, and I discussed that in my previous blog entry here.

La Brana 1 and Loschbour were both classified as part of the West European Hunter-Gatherer (WHG) mata-population by Lazaridis et al., even though only a partial sequence from La Brana 1 was available at the time. As far as I can see, the results in Olalde et al. based on the complete genome don't contradict this classification, because they show that La Brana 1 is most similar to present-day Europeans from around the Baltic Sea, just like Loschbour. Note, for instance, the position of Swedes (SE) and Poles (PL) on the far right of these graphs, indicating inflated allele sharing between them and La Brana 1 relative to other Europeans.

Unfortunately, I have to say that the main Principal Component Analysis (PCA) from the paper isn't as informative as it could have been, due to the large number of Finnish individuals included in the analysis. It's mostly a reflection of the recent population growth, founder effect and genetic drift among Finns, particularly those from eastern Finland.

Nevertheless, note that all of the non-Finnish Europeans more or less fall along the cline that runs from La Brana 1 to present-day Cypriots. This suggests that Europeans today are mostly the product of mixture, in varying degrees, between indigenous European hunter-gatherers, like La Brana 1 and Loschbour, and immigrant Neolithic farmers from the East Mediterranean. So it's a result that basically agrees with the findings of Lazaridis et al.

Interestingly, Loschbour and four other Mesolithic samples from Lazaridis et al. belonged to Y-chromosome haplogroup I, which is not at all closely related to C6. This hints at the presence of a diverse Y-chromosome gene pool in pre-Neolithic Europe, and indeed I'm still confident of seeing R1 and/or R1a among Mesolithic remains from Eastern Europe.

Even though the vast majority of haplogroup C clades are today specific to Eastern Asia, Oceania and the Americas, C6 has only been found among a handful of individuals from across Southern, Western and Central Europe, many of whom are listed at the FTDNA haplogroup C project (look for the V20+ results here). It's difficult to say when this marker or its ancestral lineage migrated to Europe, but C is one of the most basal human Y-chromosome clades, so it could represent the very first Anatomically Modern Human (AMH) wave into Europe, which actually isn't a new concept (see Scozzari et al. 2012).

The Olalde et al. paper includes a lot more information than I'm willing to cover in this blog entry. If you don't have access to the main report, please note that the extended and supplementary data are very detailed and open access.


Olalde et al., Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European, Nature (2014), doi:10.1038/nature12960


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Matt said...

Looking only off viewing the supplements and extended figures.

The f3 statistics (with Yoruba as outgroup) at look to confirm Lazaridis findings with respect to their population tree model.

The Karitiana-f3 statistic is noticeably higher in MA-1 than in La Brana (and present day Europeans), while the Han-f3 statistic is essentially identical, and within present day European variation. MA-1 seems as close to Karitiana as what are presumably Uyghur/Hazara samples, while being only as close to Han as modern Europeans.

(Interesting to note that their world PCA in the supplement at places La Brana at the terminus of a (Finnish?) European cline leading towards Han, but that is not the case for f3 statistics.).

Also interestingly does seem like some phenomenon whereby their Middle Eastern samples have a low excess of closer f3 statistics to La Brana than MA-1. It's a shame these samples haven't been labelled. The Oceanian samples and East Asian samples may have a slightly higher affinity to Mal'ta than La Brana, but it does not look very obvious.

On the D statistics, the significant findings here seem to be -

- Mal'ta is significantly closer to La Brana than Mal'ta is to present day Southern Europeans, but La Brana is not closer to Mal'tathan the two Northern European (Orcadian and Russian) samples. Presumably a combination of fairly high La Brana related ancestry in Northern Europeans, plus more MA-1 related ancestry in Northern Europeans than La Brana.
- Mal'ta is significantly closer to La Brana than Mal'ta is to East Asians
- Australian Aborigines are no closer to La Brana than they are to present day Europeans
- Aborigines are significantly closer to East Asians than they are to La Brana

Davidski said...

Thanks for the summary, I'm still getting my head around these results. Fascinating stuff.

Grey said...

(Disclaimer: This is just me idly speculating.)

Taking a simplified LGM ecozone map and Laziridis' population groups

1) West Eurasian could represent a latitudinal block from Iberia/NW Africa as far east as Central Asia/India(?) with WHG being the far western portion.
2) ANE could represent an originally more northerly population which survived the LGM on the mammoth steppe.

If so then

If ANE was derived from one of the more southerly populations then

Option 2a) ANE was derived from WHG
Option 2b) ANE was derived from West Eurasian before it split into WHG and EHG
Option 2c) ANE was derived from the EHG part of West Eurasian
Option 2d) ANE was derived from a separate East Eurasian forager population

If La Brana is WHG then it should be discard some of these options.


"But perhaps this isn't the full story, and present-day European pigmentation traits are also sourced from a late migration into Europe of a prevailingly blond people from somewhere in what is now Russia?"

I don't think that's far-fetched. If the SLC versions of lighter-skin gene started around some of the early high-density forager or farmer sites e.g. Jericho or Eastern Anatolia, and if the IE developed from foragers around the Black Sea then the lighter skin genes started much closer to them than western Europe so it wouldn't be surprising if they got them earlier.

Even if it started in India the Black Sea region could easily have got it before Western Europe.


"Even though the vast majority of haplogroup C clades are today specific to Eastern Asia, Oceania and the Americas, C6 has only been found among a handful of individuals from across Southern, Western and Central Europe"

Could that be survivor bias? i.e. various forms of C were all over Eurasia at one point until they were swamped by later adaptions but they were swamped more in some regions than others?

Grey said...

"The Karitiana-f3 statistic is noticeably higher in MA-1 than in La Brana (and present day Europeans), while the Han-f3 statistic is essentially identical, and within present day European variation. MA-1 seems as close to Karitiana as what are presumably Uyghur/Hazara samples, while being only as close to Han as modern Europeans."

(wild speculation disclaimer)

In *and* Out of America with two bottlenecks, one in each direction?

Grey said...

so in the above EEF would be one or more of
-Basal + West Eurasian
-Basal + WHG part of West Eurasian
-Basal + EHG part of West Eurasian


Basal more neo-Basal i.e. derived from West Eurasian and only seeming Basal because so common after the spread of farming?

Davidski said...

I'm still leaning to EEF being Basal Eurasian from the Arabian Peninsula, and in fact East African like and linked to hg E, something WHG-like from the Near East, maybe linked to hg J, and WHG-like from southern Europe, essentially La Brana-like. Or is that what you mean?

So a complex mix, and hard to pin down without ancient genomes from the Balkans and the Near East, and even Africa.

About Time said...

Just because Basal was maybe living in East Africa (and maybe mixed into some East Africans since then) does not mean it was equivalent to Somalis, Ethiopians, etc. (but maybe). Maybe Basal was originally more like "Ethiopian/Somali minus SSA" (more or less), and yes hg E would be a possibility.

My running concept of Basal (pre-pre-Natufians is the theory) is still that they were descended specifically from the portion of the OOA that did not become hunters. I suspect this maybe has an evolutionary component too: they were cognitively averse to megafauna hunting (just as many people are today) or the social organization in the hunting bands (too strict?). Plus they could have simply been afraid of the Neanderthals.

If that sounds too speculative, remember that Africa is the only place that nobody really hunted the megafauna to extinction and no animal domestication took place. That's all the OOA hunter group that did those things, due to adaptive differences in culture or even neurophysiology (difference in pain reactions, reactions to fear, adherence to procedure of hunting band, etc).

These peaceful non-joiners preferred to stay near the origin of the range expansion and live life as they chose (sounds too anthropomorphic or modern? If so, it's because they were "us" and not too different psychologically from us). In other words, I think of the Basals as sort of 1960's Hippies/peaceniks who hung around and "did their own thing," which became an evolutionary segue to sedentism and later - agriculture :-) Which as it turns out, lots of other people liked and copied later on. Something negative became something positive in our evolutionary history.

Grey said...

Yes, although i confuse myself on this regularly. of the logical options based on Lazaridis I think one of them would be

"West Eurasian" covering a very wide area with WHG as the westernmost branch and a more easterly EHG branch.

The missing EHG branch related to WHG (e.g. I to J just for example)

The missing EHG branch mostly doesn't exist in its original form because it mixed with Basal (e.g. Arabia, Egypt or somewhere like that) to become EEF

(also assuming this EEF later mixed a bit with WHG in Europe as well to confuse things more).

So this would mean the Lazaridis diagram should have an EHG bubble under West Eurasian with an arrow to EEF.

Grey said...

(wild speculation disclaimer)

or even the other way around, EHG -> first farmers (Sumer etc) and "Basal" came out of Arabia and squished them to form EEF.

Krefter said...

Davidski it is now called Y DNA C1a2-V20 it has relatives in Asia.

Davidski, before you said we should be cautious about the pigmentation results and now it seems you are assuming La Brana-1 and Loschbour were a deep shade of brown. I consider other possibilities and see the possibility he had light skin(there is some good evidence) a lot more plausible than most. People make the assumption that light skin mutations in gene's SLC24A5, SLC45A2, and TYR=super white skinned European. When now know through ancient DNA these are near eastern not European mutations.

I encourage everyone to read my thread Did Mesolithic Europeans have dark skin, dark hair, and blue eyes?

The light skin mutation in gene SLC24A5 is as popular in Europeans as it is in west Asians. The light skin mutation in gene SLC45A2 is more popular in Europeans though but doesn't seem to be by much. The light skin mutation in gene TYR is just as popular in Europeans as in near easterns.

Most Europeans trace most of their ancestry to native European hunter gatherers(like La Brana-1) and Stuttgart like farmers(and other types of near eastern ancestry). We know that Loschbour ~8,000 year old hunter gatherer from Luxembourg did not have any of the three light skin mutations. I couldn't pay the money to read La Brana-1's paper but I did see on the review it said he was missing the light skin mutations in genes SLC24A5 and SLC45A2(the most popular ones).

Stuttgart an ~7,500 year old farmer from Germany had alleles C/A in SNP rs1042602 according to SNPedia is about 50% in Europeans and is the light skin form. In SNP rs16891982 in gene SLC45A2 she had alleles C/C so the dark skin form. In SNP rs1426654 n gene SLC24A5 she had alleles G/G so the light skin form. Those results are typical for any modern near eastern or European. Except it seems the one in gene SLC45A2 is more popular in Europe. I have heard Otzi an ~5,300 year old farmer form the alps has the same results as Stuttgart.

Isn't this good enough evidence to say the light skin mutations were brought to Europe with near eastern like farmers? I think these mutations are a sign of near eastern not European ancestry. Many have made the mistake to only connect these mutations to Europe.

How do you explain paler skin in north Europe than in south Europe and the paler skin in southern Europe than in the near east even though they all have around the same amount of these light skin mutations? Obviously there are other factors to European pale skin(especially in northern Europe).

Today farmer in Europe farmer ancestry correlates with olive skin, dark hair, and dark eyes(highest in southern Europe) while hunter gatherer ancestry is the opposite it correlates with light skin, light hair, and light eyes(highest in northern Europe).

A lot of blue eyes in a population means there is some light hair and very pale skin, that is just fact. People ignore that fact because they assume Loschbour and La Brana-1 had dark skin. Their argument towards me are random pictures of blue eyes Indians. What percentage of Indians have blue eyes? Argument is over.

Is there some way to get Melanie or whatever from La Brana-1 to actually see what skin color he had? I think he most likely had pale skin. For the reasons I named above. One big question to ask is why are Europeans who have the most Mesolithic ancestry also have the palest skin?

Krefter said...

Can you guys please argue with me? Everyone seems to assume these mutations always cause light skin and I just gave good evidence that is not the case. Most of the time people ignore me and keep assuming those mutations are European and always cause pale skin. I am saying we don't know what skin color La Brana-1 and Loschbour had it very well could have been dark or light.

Krefter said...

It seems La Brana-1 had a lot of ANE ancestry. The origin of ANE in Europe and the near east is more complicated than Indo Europeans and eastern Europe.

About Time said...

@Barak, my money is on light skin as an adaptation to Vitamin D deficiency for farmers. Could have started in Europe or any cloudy area (in Gedrosia territory? Eastern part of SLC24A5 mutation map looks exactly like Gedrosia).

Stuttgart had light skin. The European hunter/gatherers so far had dark skin/hair and light eyes. Malta had dark skin/hair/eyes.

I keep asking the same question: Why would hunter/gatherers suddenly become light pigmented (blond/blue/fair skin) if they were dark pigmented eating the same diet and living in the same environment? It makes no sense in Darwinian terms.

Farmers on the other hand, would be under environmental stress from bad diet. They would also (IMO) be living in tight knit communities with inbreeding depression but also the evolutionary chance to fix beneficial alleles. I'm assuming EEF were "leapfrog" colonizing areas as intact communities and not mixing with hunter-gatherers, just as Jamestown colonists didn't mix with Amerinds.

Mixed communities would be the fringe areas around EEF settlements, where WHG could leverage their animal handling skills to manage livestock. This segued to a new specialization: pastoralism.

We know about EEF from Stuttgart, but it also mixed into Arabs, Caucasus, Uzbeks and Gujarat. Origin of EEF could be anywhere. Kurgans could have been partly EEF, partly ANE. Thus light pigment in Andronovo etc could be EEF origin too.

Again, I think that because why else would Siberian hunter gatherer descendants get light hair/eyes/skin, when we know Mal'ta was dark pigmented. There is no evolutionary reason for it, unless they mixed with light population (EEF) and then selected those alleles for any reason (sexual selection or even kinship marker between related pastoral clans that moved around, etc). That would at least make Darwinian (i.e., objective scientific) sense.

About Time said...

Sorry to double post, but I keep see the same points being made. A lot of logic people bring up implicitly assumes that because we see several traits together today (like light hair + light eyes + light skin), those traits were always assorted that way.

That is, basically, "Typological" thinking. Again, that's way outdated. It's maybe something we all are predisposed to do (it's a cognitive shortcut), but it's not scientific and its specifically not Darwinian.

Law of Independent Assortment.

The Law of Independent Assortment, also known as "Inheritance Law", states that separate genes for separate traits are passed independently of one another from parents to offspring. That is, the biological selection of a particular gene in the gene pair for one trait to be passed to the offspring has nothing to do with the selection of the gene for any other trait. More precisely, the law states that alleles of different genes assort independently of one another during gamete formation. While Mendel's experiments with mixing one trait always resulted in a 3:1 ratio (Fig. 1) between dominant and recessive phenotypes, his experiments with mixing two traits (dihybrid cross) showed 9:3:3:1 ratios (Fig. 2). But the 9:3:3:1 table shows that each of the two genes is independently inherited with a 3:1 phenotypic ratio. Mendel concluded that different traits are inherited independently of each other, so that there is no relation, for example, between a cat's color and tail length. This is actually only true for genes that are not linked to each other.

Europeans are are convergence of at least three populations (maybe with at three evolutionary histories - especially EEF vs WHG/ANE). For all we know, blondism arose from small gracile EEF farmers living in a cloudy area with Rickets ( or similar health problems. Evolutionary pressure was compounded by living in crowded farm settlements with non-optimal water and sanitation, livestock, etc.

Some EEF mixed with brawny and dark hair/skin blue eyed WHG. Some EEF+WHG descendants would have any combination of these or any other traits. Some could be blond haired and blue eyed, with better resistance to European pathogens from La Brana (just an example).

If groups were then migrating off in different directions, these traits could randomly diverge or be sexually selected for (just like in any organism). In time the distributions of these originally independent traits could be similar. We would then (wrongly) assume they were always part of a "package of traits" or "type."

Fanty said...

I wonder if there still is the possibility of a light skin allele on their own?
I mean, East Asians have white skin based on their own (indepentand of the European/SW Asian/S Asian) one too?

So, if they lack a Caucasian white gene and a mongolid white gene, they still may have their own white gene? *scratch head*

Different thing...
I am wondering on the 180 degrees turn in food quality recently. Some years ago, it was commonly claimed farmers had the food producing advantage over everyone else and thats why they outbreed all other races.

And now (since a few month only or so) more and more scientists claim, Farmers had a bad diet, they didnt get as much food as HG, they also didnt get as good quality food as HG. They suffered from a lot food based illnesses that deformed their skeletons and hindered their overall development ....and they even had a shorter life expectancy than HG.

HG on the other side are "healthy", lived long timespans, never had any famine as much as their bones tell and are generally well developed physically from very high quality food in huge amounts...

Oh... really? All of the sudden, the praised "one and only super successfull lifestyle" is rated as "total crap". Why would HGs then adapt farmer lifestyles at all, if its so bad?

Grey said...


"Can you guys please argue with me?"

I'm reading up on skin genes :)

Apparently there are seven known: the 2 x SLC ones, KITLG, AGIP, OCA2 and umm, haven't got that far yet, and La Brana had 3 of them (or 2 1/2) out of 7

It might be possible to calculate it exactly by adding and subtracting which ones La Brana had and how big of an effect each one has but i don't know enough to say.

Personally (cos of the whole mammoth steppe thing) in terms of shade i think they'd be in the same ball park as Native Americans (or like very tanned white people) but i don't know enough to say for sure.

I think the skin color genes originally started in the mid-latitudes and spread because they had some other beneficial effect with skin lightness as a side effect. *However* they do seem to have been particularly strongly selected for in Europe and especially in the north and northwest of Europe so i'd guess either there was secondary selection on skin color in itself *or* the beneficial effect was more critical in that region for some reason.

the links i'm reading!&

Grey said...

"One big question to ask is why are Europeans who have the most Mesolithic ancestry also have the palest skin?"

It might simply be a double correlation with the region i.e.
-people in that region had the strongest selection on skin color
-WHG in that region survived better in that region than further south

Are the hap I people in the south of Europe light or dark? (assuming hap I is the main WHG group)

Krefter said...

About Time,

There are a gazillion possibilities out there. I don't think anyone fully understands how evolution happens a lot of it seems impossible to explain but we know it happens. It is almost like Evolution is a living thinking thing inside of us. It mutates in our descendants or something.

Did you read my post and like a million other posts I made about the same subject?

Those three mutations DONT CAUSE WHITE SKIN. They are about as popular in near easterns as they are in Europeans. There are other factors to European light skin that have not been discovered. They are near eastern NOT EUROPEAN MUTATIONS. Europeans have them because of their near eastern ancestry.

I think there is a high possibility that La Brana-1 and Loschbour had light skin just we don't know what mutations cause it yet.

People always assume the farmers were the light ones. Today farmer ancestry in Europe correlates with dark hair, dark eyes, and olive skin. Hunter gatherer ancestry correlates with light hair, light eyes, and light skin. The farmers mainly descended from the same source as modern near easterns do who we know are very dark. Hunter gatherers being dark MAKES NO SENSE. People can't find an explanation for that.

You have to create some very crazy theories on the rapid evolution within about 3,000 years that made all of Europe go from dark brown to white!!! Dude, do you see how crazy that is? Obviously it is a lot more complicated that that.

I think it is more likely that some Mesolithic Europeans were dark and some were light and it happens that the light ones are the main ancestors of light Europeans. So it seems hunter gatherer ancestry correlates with lightness. I think it is likely the eastern European hunter gatherers were the light ones. The reason I say that is because of Indo Europeans and Uralics. There is evidence the lightest Europeans trace most of their ancestry to Indo Europeans or Uralics from eastern Europe. We also know the people who spread Indo Iranian languages were mainly light haired and eyed.

MA1 lived 24,000 years ago!! No duh he was dark. Also they don't know what skin color he had all they know is he did not have the three light skin mutations. Which I have already shown are not the only factors to creating pale skin in Europe.

It is debatable how much ANE ancestry the bronze and iron age Indo Iranians had. They had over 30% mtDNA U2e, U5, and U4 all typical of Mesolithic Europeans(but of course people will ignore that). Also light hair and eyes today correlate with Mesolithic ancestry not ANE.

I guess it is possible there was strong selection in Europe during the Neolithic and that pigmentation features in different areas of Europe have different origins. Is till think it is very unlikely la Brana-1 and Loschbour had light skin.

Grey said...

Apparently there are 7 skin lightening genes and La Brana had 2 1/2 or 3 out of the 7


"Why would HGs then adapt farmer lifestyles at all, if its so bad?"

quantity over quality

100 weedy farmers vs 10 super-fit HGs -> HGs still lose
100 weedy farmers vs 30 semi-fit forager/herders -> ?

Grey said...

"Those three mutations DONT CAUSE WHITE SKIN."

There are seven (apparently). Some have more effect than others and each one can go from 0% in a population to 100%. They add up.

About Time said...

@Fanty, it's not that simple. Conditions can be in some ways unhealthy for the individual organism but evolutionary advantageous for the population. F.e. if the "bad" conditions help select for beneficial alleles and support population growth (sometimes in spite of poor health, etc). That's exactly how Darwinian processes work. "Spew and skew."

HGs might have had an OK lifestyle, but if it wasn't changing and wasn't selecting for new beneficial alleles, it was stagnant and in the long run less advantageous than the "unhealthy and cramped and inbred" farmer settlements. Especially if the ecological resources were slowly dwindling.

Also, I suspect the farmers were better socially organized, with a more rule bound society (necessary to regulate planting/harvesting times, etc). That has its own advantages. But the hunter might ask why those farmers are doing all that bizarre seed counting or digging in the dirt when they could be relaxing, etc (deferred gratification).

So one day the relatively healthy hunter can laugh at the strange and apparently "unfit" farmer, but then 1000 years later it might be the farmer that has more descendants. Or in Europe, it looks like the mix of EEF plus WHG left descendants in most places in the long run.

Krefter said...

What are these 7 light skin genes(mutations)? The east Asian one I have heard of is centered in southeastern Asia not in Siberians who have the same skin pigmentation.

About Time said...

"100 weedy farmers vs 30 semi-fit forager/herders -> ?"

@Grey, don't forget heterosis. If the EEF had some squirrely deleterious mutations (inbreeding depression), most those would be absent in the first generation of mixed EEF and WHG and less pronounced in later generations.

Mixing would get a chance to select for the best of both worlds (optimize all beneficial traits). So you potentially could have 30 super fit mixed pastoralists, with more brains than the foragers and more brawn than the farmers. Courage of a hunter with meticulousness of a seed counter. Etc.

Grey said...

@About Time

Yes, true. Either way, quantity has a quality all its own :)


i'm still reading those links i posted: there's the two SLC ones, TYR, KITLG, OCA2, AGIP and umm missing one.

Anonymous said...

"So as things stand, it appears that Europeans only acquired their present coloring, including pale skin and a high incidence of light eyes, relatively recently, well after the hunter-gatherers and farmers began mixing"

We can't still be sure of that. So far all these mesolithic individuals were "out of place" haplogroup-wise ('I'm referring to the I2 individuals of Sweden and the one from Luxembourg too IIRC).
It looks like somehow they've been replaced, including by other I, like I1 (that we can bet were hunter-gatherers in Europe too, somewhere else).
Since the I1 are among the popoulations having the lightest skin and hair and eyes, we should wait to have several samples of mesolithic aDNA with I1 to be sure.

Besides the fact that these C6/I/I2 had sometimes blue eyes might be a clue that some European HGs were carriers of light features somewhere else, and that by non-frequent admixture events, it entered these sampled mesolithic populations, and that the ancestral population (yDNA I) of HGs that entered Europe had the blue eyes' mutation and that some particular population (ending up as I1) became lighter, (the R1a getting these characteristics by mixing w/ them) or maybe R1a1a was indeed really the source of lighter features - but it seems the blue eye's mutations was already present in non-R1a mesolithic HGs anyway - before the R1a got wester).

Grey said...


Fanty said...

"It looks like somehow they've been replaced, including by other I, like I1 (that we can bet were hunter-gatherers in Europe too, somewhere else)."

The MRCA of I1 is said to have lived 5K years ago.
Well, that doesnt mean, there had not been I1 before that time, but ALL MODERN I1 stem from a single male that lived like 5K years ago.

those I2 whg-guys had been 8K years old. 3K years older than the common anchestor of all modern I1.

Finding mesolithic I1 would indeed be something special as this would be before the 1 person bottle neck that the I1 "people" went through at 3000BC.

Anonymous said...

Fanty: "The MRCA of I1 is said to have lived 5K years ago."

I'm not a big fan of this kind of estimate, so, as far as I'm concerned, I1 or at least this I1 individual you're talking of, might be older.

pyromatic said...


Why? What is your age estimate? What method do you use to arrive at it? Are you not a fan simply because it contradicts some belief you may hold?

Grey said...

"The MRCA of I1 is said to have lived 5K years ago."

Around the time of Funnelbeaker.

"Finding mesolithic I1 would indeed be something special as this would be before the 1 person bottle neck that the I1 "people" went through at 3000BC."

If it's Funnelbeaker and it started at the edge of the LBK and/or megalith farmers and spread north and east towards the Baltic then that's where to look imo - around the edge of the Baltic.

Krefter said...

The Swedish funnel beaker farmers were basically the same as Stuttgart and Otzi so I doubt much changed as the farmers went north. The reason I think modern north Europeans have so much WHG is because of migrations in the metal ages by Indo Europeans and Uralics.

"Are the hap I people in the south of Europe light or dark? (assuming hap I is the main WHG group)"

Well y DNA isn't everything what type of results do they get in autosomal DNA? Much less WHG than most northern Europeans.

Chad said...

I think we need to forget about blonde, blue and pale going together. They must've started independently. Just because a section of the world has all three together doesn't mean they go together. We can have a blonde, brown-eyed Mongolian, a pale, blue-eyed, dark haired Brit. We can have a light skin, dark hair and eyed Near Easterner. Sexual selection is looking like a big factor in at least the hair and eyes.

Skin may be more of the adaptation to the diet. I say that because we still see dark complexion in Siberia and America, probably because they kept the fish and meat diet. The cline in Europe to the Near East can easily be explained by the later arrivals from the Near East, probably containing East and Sub-Saharan admixture.

I believe that most Europeans show the same reflective qualities in the underarm area, regardless of how dark the rest may look. It is just that some live in a climate where there is more exposure.

I1 may make an appearance in the East Baltic or Scandinavia with the Pitted Ware. It started about 2800BCE so the MRCA of 5kya would fit nicely.

Matt said...

Beleza (2013) found that in a mixed Cape Verdean population, % European ancestry explained as large again an influence on skin color independently than the sum of the "four major loci SLC24A5... TYR... APBA2... SLC45A2"...

"that together account for 35% of the total variance, but the genetic component with the largest effect (~44%) is average genomic ancestry. Our results suggest that adjacent cis-acting regulatory loci for OCA2 explain the relationship between skin and eye color, and point to an underlying genetic architecture in which several genes of moderate effect act together with many genes of small effect to explain ~70% of the estimated heritability."

(in pie chart form -

The supplement to Olalde seems right to say "The existence of many association studies on pigmentation and the understanding of the biochemical pathways involved in the melanosomes indicate that any new discovery on gene pigmentation will represent a very minor proportion of the observed phenotypic variation." Any new correlated individual gene variation will represent very little.

But there may be many, many genes of small effect that we currently don't know about. And there may be a lot more space in terms of the existing known genes explaining less than we've thought - some of the previous estimates may not have controlled sufficiently well for ancestry (the Cape Verde population studied by Beleza might have more linkage disequilibrium between the genes in question and overall ancestry, allowing better estimates of their power, compared to previous African American studies).

It might be that light skin is a side effect of selection for... something else... but it would seem strange to have many genes of small effect pointing in that direction if so. Not inexplicable, because there could be many genes which either produce pigment or do something else useful, and once the selection for protective dark pigment goes away (not much sun, people have more clothing, etc.) they all uniformly get selected to do something else useful (doing the same useful thing seems unlikely though).

Anyway, although the estimated dark skin is not to be dismissed, there may be some hidden heritability of skin colour which might give La Brana and Stuttgart a lighter pigmentation than we would predict (especially as in some ways they have more "European" genomic ancestry than any present day population). No real reason to bet on it, but there's some room there.

About Time said...

Would be good to see pigment genes for Gok4. It could give at least one data point to test whether my "blond EEF population" theory holds up, especially bc I think the "blond EEF" were precisely Funnelbeaker.

Grey said...

What i want to find out is what skin-lightening genes Native Americans have compared to La Brana.

Grey said...

mixture of light and dark and dark and light

Grey said...

"The Swedish funnel beaker farmers were basically the same as Stuttgart and Otzi so I doubt much changed as the farmers went north."

If you look at the Funnelbeaker distribution and assume the process of creating a shifting agriculture started along the borders of LBK in the south and/or megalith culture farmers in the west *and* if that culture then spread north and east towards the Baltic then it would have started as mostly EEF around the edge and became more WHG the further it went from the start point towards the Baltic coast.


Fanty said...

I painted and lumped together new maps with WHG mtDNA, y-DNA, eye color, phenotype regions etc.

Grey said...


Yes to your main point. I don't know enough to say.

"It might be that light skin is a side effect of selection for... something else... but it would seem strange to have many genes of small effect pointing in that direction if so."

I was wondering if halfway between light and dark skin might result simply from removing the selection for dark skin once out of the tropics as a random positive mutation that reduced melanin might be net negative in the tropics but positive outside.

Fanty said...

the interesting thing is... in the very northern end of Scandinavia/Finland there ARE darker skinned people, but they are also black haired and brown eyed. So, they dont match the color combination observed in WHGs, even if their mtDNA is like 40% WHG/ANE like.

Grey said...

awesome maps

About Time said...

@Grey, is that a guess or from genotype?

Grey said...

@About Time
just thinking aloud. i had a random thought once that blonde might be the result of an odd mixture but i was thinking East Asian from the north as the catalyst. not based on any more than that.

Grey said...

My understanding, possibly wrong

West Eurasian

West and East Eurasian

SLC24A5 and KITLG between them are about 50% of lightness and La Brana didn't have them.
SLC24A2 is associated with olive skin. I don't know if that means it is a lightening gene in itself or it just modifies others but La Brana didn't have it either.

I think that means La Brana had

Other points
1) OCA2 is associated with skin lightening in East Asians but blue eyes in Europeans
2) ASIP and KITLG are old and it's odd that La Brana doesn't have KITLG
3) I think the two SLC ones are the recent (agriculture time) ones
4) Apparently there are cross-effects from eye and hair color i.e. the way the various skin lightening alleles display might vary depending on eye and hair color genes.

map of distribution of five of them but not KITLG

(plus neanderthal genes - not sure if that is relevant or not)

Krefter said...

I know many of you are probably sick of me talking about pigmentation because Y DNA, mtDNA, and autosomal DNA are much more important. But I just found some interesting stuff, Stuttgart for sure had brown eyes(but actually maybe her eye color is unknown) and Loschbour for sure had blue or green eyes and it seems both had dark skin according to the 8-plex system(predicts skin and eye color). Of course the 8-plex system could be inaccurate.

In Laz 2013 the SNP's they had a list of SNP's associated with skin and eye color under the section 8plex SNP's for hair color it was under Hiriplex SNP's.

I found the study that created the 8plex system. It was made for the police but doesn't make it any less effective for 7,500 and 8,000 year old DNA. Below is a link to the study.

And below is another link to the system they have to predict eye color.

Here is an description of the predictor.
"The prediction for eye color is a two-step procedure: First, rs12913832 (HERC2) distinguishes eye colors as being not blue (ie, brown or green) or not brown (ie, green or blue). Further distinctions for predicting eye color as being brown, green, or blue are dependent on the genotypes of three additional SNPs: rs12203592, rs16891982, and rs6119471 (Figure 2). Brown eye color is predicted by the following genotype combinations: A/A or G/A at rs12913832, plus either G/G at rs6119471, or C/C at rs16891982; green eye color is predicted by G/G at rs12913832 plus C/C at rs16891982, or by G/A at rs12913832 plus T/T at rs12203592; and blue eye color is predicted by G/G at rs12913832 plus T/T at rs12203592.

The second step proceeds with samples that were not positively described as being brown, green, or blue in the first step (Figure 2). Samples that are homozygous for G/G at rs12913832 (ie, not brown) plus T/T at rs12896399 are then predicted to have blue eyes, while samples that are homozygous for A/A at rs12913832 (ie, not blue) plus G/G at rs12896399 are predicted to have brown eyes. This step utilizes the newly-included SNP rs12896399 (SLC24A4) and leads to increased numbers of positively described eye colors."

Eye color test

Loschbour: rs12913832 G/G(blue or green), rs12203592 T/T(blue), rs16891982 C/C(green). They never specified whether green eyed people with rs12913832 G/G don't also have rs12203592 T/T. All they said was that green eyed people with rs12913832 G/A have rs12203592 T/T. So I guess Loschbour according to this system had blue or green eyes.

Stuttgart: rs12913832 A/A(brown or green), rs6119471 C/C(not positive for brown eyes), rs16891982 C/C(brown eyes). They said that brown eyes is determined in step one by rs12913832 A/A or rs12913832 A/G and either rs6119471 G/G or rs168982 C/C. I am not sure if the combination has to be rs12913832 A/A rs6119471 C/C and rs12913832 A/G rs16891882 C/C. If it is that means Stuttgart's eye color is unknown because she doesn't fit blue or green eyes. But I doubt it has to be fit that way which means Stuttgart had brown eyes.

The error rate they said for eye color prediction was 5%. They also admitted that oversampling non Europeans because all humans except Europeans are practically 100% brown eyed. 69%(55/803) of the people they tested for the eye color test were European and 17.8%(143/803) were described as mixed and showed some non brown eyes. So hopefully this means it can accurately predict eye color. It was also said that the errors were all green eyed people who were predicted as blue or brown eyed. They said the error rate for Europeans at the second step was only 2%.

Krefter said...

The study says that light or medium skin is predicted by any is predicted by any two of the following alleles: G/G at rs12913832, G/G at rs16891982, A/A at rs1426654, T/T at rs1545397, or A/A at rs885479. I think that means if you have one of the SNP's you have light or medium skin.

They also said that Light skin color is predicted by more stringent conditions: G/G at rs12913832, plus G/G at rs16891982, and A/A at rs1426654. Non-light skin color (ie, medium or dark) is predicted by G/G at rs6119471

Light or medium skin test

Loschbour: rs12913832 G/G(light or medium), rs16891982 C/C( not light or medium), rs1426654 G/G(not light or medium), rs1545397 A/A(not light or medium), rs885479 G/G(not light or medium). Loschbour was not light or medium

Stuttgart: rs12913832 A/A(not light or medium), rs16891982 C/C(not light or medium), rs1426654 A/A(light or medium), rs1545397 A/A(not light or medium), rs885479 G/G(not light or medium). Stuttgart was not light or medium.

light skin test
Loschbour: rs12913832 G/G(light), rs16891982 C/C(not light), rs1426654 G/G(not light). Loschbour was not light.

Stuttgart: rs12913832 A/A(not light), rs16891982 C/C(not light), rs1426654 A/A(light). Stuttgart was not light.

Both Stuttgart and Loschbour had derived alleles C/C in SNP rs6119471. Ancestral alleles G/G they said cause dark skin.

For the skin color predictions out of 600 tested only 4 were errors so 1%. The 4 were dark skinned people who were predicted as light skinned. They also only tested Europeans, African Americans, east Asians, south Asians, and mix(what ever that is). They also had a bunch of inconclusive including for Europeans, and only Europeans had a high amount of not dark.

East Asians should not work for this test since their light skin is not from the same source as European light skin. They should test other west Eurasians besides Europeans because I am sure they would mostly be predicted as light skinned even though they are not.

On Wikipedia says there are three SNP's with mutations that are associated with European light skin. The first is rs1426654 A/A(in gene SLC24A5) which is close to 100% in west Asia and Europe and very popular in north Africa south Asia. Stuttgart had rs1426654 A/A. The second one is rs16891982 G/G(in gene SLC45A2) it seems to be very dominate in Europe close maybe around 80-90% and around 50% in the near east and a little less than that in north Africa and even less in south Asia. Stuttgart had rs16891982 C/C. The third light skin mutation is rs1042602 C/C(in gene TYR) it seems to range in Europe 50-25% and the same for the near east but it is less popular in south asia. According to SNPedia around 70-80% of Europeans have either C/C or A/C. Stuttgart had A/C.

Based on that you cant say Stuttgart had light skin. He has the same alleles as probably most modern near easterns. It is very important to not he did not have rs16891982 G/G which is the only one that is significantly more popular in Europeans than near easterns. Based on Stuttgart's closest modern relatives(Sardinians) I think she had light skin possibly olive or brownish.

Shockingly if you go by 8-plex Loschbour had blue eyes(maybe green but cant confirm) and not light or medium skin and not dark skin. I don't know exactly what the dark skin thing means or how it is distributed in the world.

I cant pay for the study on La Brana-1 but they say he had blue eyes and I was able to read in a preview that he was did not have rs1426654 A/A and rs16891982 C/C the two that are close to 100% in modern Europeans. Of course it is important to remember the only one that is more popular in Europeans than in west Asians is rs16891982 C/C.

Dark haired, dark skinned, and blue eyes it is hard to believe that was a common phenotype in Mesolithic west Europe. I still think there is a possibility they didn't have blue eyes or they didn't have dark skin.

Krefter said...

Maybe many Mesolithic Europeans looked like this. Does anyone remember the movie 10,000BC? It was about hunter gatherers from the Urals. They had dark skin, dark hair, and there was one woman known as the blue eyed girl. Here people were killed and she was adopted.,d.aWM&psig=AFQjCNE2RVs-v-S-j_-ywVOrJWcivJKbYA&ust=1390967742531689,d.aWM&psig=AFQjCNFP2_zO5PBdbUuoYP_GKeW9urcg_Q&ust=1390967905363063,d.aWM&psig=AFQjCNFP2_zO5PBdbUuoYP_GKeW9urcg_Q&ust=1390967905363063

Fanty said...

Well, since the modern distribution of blue eyes does very well fit the WHG like autosomal DNA, I would say, the tests have it right, blue eyes must have been common in WHG populations.

But I cant aswell, imagine blue eyes and dark skin to be something "normal".
I cant even look at such people, they scare me to
It looks totaly "wrong".

*shudders*..... its THEM.... the white...err black walkers!

Fanty said...

I recall that movie.
The problem is that SHE is white. But isnt she suposed to be the first blue eyed human ever born? (and the mother of all the modern blue eyed humans etc) ...then it seems she needs to be black. X-D

I think the movie was inspired by the Danish paper that said, the blue eyes did come up in a single human and all modern blue eyed people can trace back their anchestry to that single human.

Krefter said...

Thank you very much for the maps Fanty. You just guessed what how WHG ancestry many regions had right? Because Not every little region in Europe has been tested but you could make accurate guesses based on what people close to them had.

I have seen the same light eye map from the 1960's. It seems back in the 1800's and early 1900's people were much more interested in Anthropology than today. The reason may be because your seen as raciest if you are really into it. It is true people who are into anthropology can be raciest and especially back then(Nazi ideology had a lot to do with Anthropology).

I think those "crazy raciest" from the 1800's and 1900's have a lot of info that we can use as a tool with DNA. I am shocked by how similar that map is of WHG like skulls with light hair, light eyes, light skin, and Mesolithic European ancestry. Obviously those guys in the 1940's knew what they were talking about. Now some 70 years later it has been proven Mesolithic European ancestry is distributed in a very similar way.

I doubt the ANE+WHG mtDNA map is accurate. mtDNA U5, U4, and U2e(dominate in Mesolithic Europeans) combined are not strongly centered in any area of Europe. They have a pretty even distribution for most of Europe.

Where did you get the light skin map? Southern Europeans are much lighter than near easterns and north Africans. Today I was arguing that south Europeans are significantly darker than northern. Then a Spaniard showed me like a gazillion pictures of Spaniards from every little sub region of Spain. They don't really look much different from north Europeans. Besides being almost all are dark haired and eyed and there were some with darkish skin.

If the light skin map is legit it obviously correlates with light hair, light eyes, WHG ancestry, and WHG skull shape. LIGHT EYES ARE CONNECTED WITH LIGHT HAIR AND LIGHT SKIN.

This is why before any pigmentation genes were taken from Mesolithic Europeans I thought they were nearly all light haired and eyed and also that they had very pale skin. It is honestly shocking that La Brana-1 and Loschbour had blue eyes, dark(probably black) hair and dark skin.

Krefter said...

If a Time Machine is ever invented in our lifetime NEVER GO BACK TO MESOLITHIC EUROPE!!!

Krefter said...

The actor doesn't look very white.,d.aWM&psig=AFQjCNHI0Yx2GL_CZDSAn7mrTz0R5XjD5w&ust=1390971629356456

I doubt the first person with blue eyes lived just 12,000 years ago. If it is true Loschbour and La Brana-1 had dark skin and blue eyes I doubt the first person with blue eyes was light skinned.

Krefter said...

I remember that her people were killed and she was adopted. I haven't seen the movie for a long time so I may be wrong. I think I remember seeing some people from her tribe and they all looked very white. The people didn't even have Caucasian facial features so that part is un accurate.

Fanty said...

"You just guessed what how WHG ancestry many regions had right? "

Its a blend between the original paper values and the ones from the calculator.

"I doubt the ANE+WHG mtDNA map is accurate. mtDNA U5, U4, and U2e(dominate in Mesolithic Europeans) combined are not strongly centered in any area of Europe. They have a pretty even distribution for most of Europe."

I superimposed all the U maps from Europedia in photoshop. Then I made the gradient 3 colored to better see whats the centers of dirtsibution and tried to simpyfied draw it on a blank map.

Krefter said...

Take a look at Fanty's maps and just research a little and you will learn light skin, hair, and eyes are very connected. strangely they all have a similar distribution to WHG(Mesolithic?) skull shape and WHG ancestry. The reason that is strange is because Loschbour and La Brana-1 had dark probably black hair and because it seems they had dark skin.

Here is an example of evidence light hair and eyes are connected.

Fanty said...

I watched the movie about 1 or so years after I read the article about the blue eyes. Thats why I made the connection.^^

Fanty said...

So it all stands and falls with the accuracity of the Europedia maps.

I somewhat doubt that Czechia is surounded by U and in itself empty of it however.

Fanty said...

"Where did you get the light skin map? Southern Europeans are much lighter than near easterns and north Africans."

Its the only map in the net that shows a gradient in Europe at all.

Krefter said...

Are there enough samples to make a very accurate map? mtDNA U3 is not of Mesolithic European origin. If that map is accurate it is very surprising. a North Norway Sami person tested in Laz 2013 had ~51% WHG the highest found so far also Sami have much darker hair-eyes-skin than other 30%+ WHG people. Also the Sami have about 50% mtDNA U5b1b1 and 50% V. Both are probably from bottlenecks. The Sami have no diversity of U5 and no U4 or U2e. Also mtDNA V has not been found in pre Neolithic Europeans but constantly appears at a low rate in Neolithic Europeans. The Sami scored about 29% EEF so they have farmer ancestry even though they are technically hunter gatherers.

Fanty said...

"Then I made the gradient 3 colored to better see whats the centers of dirtsibution"

That means I made it a gradient from blue over yellow to red.
Then I only drew the blue parts to the map at all and left the red and yellow ones white. Nobody could see anything at all if I woulnt limit the sight to the real hotspots.

Fanty said...

Well, if you say all my maps suck, then do your own ones.

Krefter said...

I don't think you should label it has ANE+WHG mtDNA. We don't really know exactly what ANE is there are just a bunch of theories. There is one mtDNA sample from MA1 and he had mtDNA U(apart of his own subclade). Why would to separate people have the same mtDNA haplogroups? Part of the reason why I think ANE ancestry is more complicated than most think is we can't find mtDNA that probably descends from them.

Krefter said...

"Well, if you say all my maps suck, then do your own ones."

LOL that's a good point. I might but I have a pretty good idea how their distributed in my head. If I ever did I would make separate maps for WHG, ANE, and EEF and put little circles graphs for each nation or region it was tested in.

Fanty said...

"Part of the reason why I think ANE ancestry is more complicated than most think is we can't find mtDNA that probably descends from them."

The paper says, ANE has the same mtDNA than WHG does. They form a unity mtDNA wise. Also they are autosomal DNA like relatively close related.

Its just that the WHG guys are all I2, except for that C6 guy now. And the 2 ANE samples we know of are R* and R1a

Fanty said...

" If I ever did I would make separate maps for WHG, ANE, and EEF and put little circles graphs for each nation or region it was tested in. "

I have something like that. All 3 with percentages in each country that was tested and in a different color the countries that are calculated.

But I decided against posting them. (the WHG map is the WHG map, just without the percentages)

Fanty said...

the problem with percentages is.... they are not relayable. Specially not the ones from ADMIXTURE. You CANT take them literate. So its better to watch the big picture (wihtout exact percentages). Thats why I let them out.

Krefter said...

There is really no evidence in mtDNA and Y DNA Scandinavians and north-east Europeans get their WHG from that process. How do you explain their R1a(Corded ware), R1b(Nordic bronze age), I1(post Mesolithic from mainland Europe), N1c1(eastern Asia)? If it was simple mixing with Pitted ware and Motola like people why do they have such a low percentage of mtDNA U5, U4, and U2e? Why are Finns and Sami's U5 almost entirely under U5b while it was the opposite for Mesolithic Scandinavians and Russians?

Why didn't they mix a lot with hunter gatherers as they were moving up north from central European?

Krefter said...

Those are the percentages given by the admixture. Who knows how accurate they are but that doesn't matter. Plus giving the percentages can give people a better idea of what the differences are.

We have one mtDNA sample from MA1 that's it. Think about it if many WHG people did not mix with ANE why would they have the same maternal lineages? How are WHG and ANE so related? Being hunter gatherers means nothing.

Grey said...

"I think we need to forget about blonde, blue and pale going together."

Other way round. It seems eye and hair color does tend to impact skin color but not necessarily in an absolute way.

Davidski said...


The study clearly says they're closely related genetically.

"Outgroup f3 and D statistics (16,17), using different modern reference populations, support that Mal’ta is significantly closer to La Brana 1 than to Asians or modern Europeans (Extended Data Fig. 5 and Supplementary Information). These results suggest that despite the vast geographical distance and temporal span, La Brana 1 and Mal’ta share common genetic ancestry, indicating a genetic continuity in ancient western and central Eurasia."

And what about all the mtDNA U among ancient remains from Siberia from different periods? What makes you think these lineages belonged to WHG individuals? So where were the ANE people hiding? And why do the Kurgan samples carry so much Y-hg R? Isn't that an ANE marker?

Grey said...

KITLG lightens the skin and also effects blond hair
OCA2 lightens the skin (in East Asians) and is a factor in blue eyes (in Europeans)

Thirdly there's MC1R which is connected to red hair.

Neanderthals were light too - plus lots of red hair - but they had a lot longer to adapt to the latitude.

But given so many ancient writers mentioned pale, red-headed peoples i'm wondering if La Bran was checked for that?

Davidski said...

It can get very complicated. For instance, I supposedly don't have any of the blond hair markers at 23andMe, but I do have an allele for red hair, and I've actually got reddish blond hair, which has gotten darker over the years, but was still very light when I was a teenager.

spagetiMeatball said...

David, how long do you think it is before we start seeing modern populations clusters as combinations of ancient populations, i.e., how many more samples should be sequenced?

Davidski said...

We'd need at least 10-20 from each region and period to be sure of anything. So maybe ten or more from Upper Paleolithic South Siberia, another 10-20 from Mesolithic Western Europe, and so on. That would make the ADMIXTURE analyses much more interesting, but also the formal mixture tests more secure.

Unknown said...

I swear someone is either slow or a troll!!!

Unknown said...

Will the whg component be split into loschbour, la Brana, and motala? Is there a way to pick out subtle differences like with Atlantic, North Sea, and Baltic this far back?

Davidski said...

If there's enough drift between hunter-gatherer populations from different parts of Europe than that should be possible. But it seems that back then Eurasia was sparsely populated and people were highly mobile, that's probably why there's such strong genetic affinity between La Brana and MA-1. So I don't think it'll be all that easy to place the ancient samples within modern country borders or anything like that.

Krefter said...

I cant read the La Brana-1 paper.

Obviously La Brana-1 had some ANE ancestry. I couldn't find any evidence in Laz 2013 WHG and ANE are closely related. They did say that EEF and WHG are closely related when you compare them to people around the world.

mtDNA U is a big and very old haplogroup not every single little mtDNA U sample is connected with European hunter gatherers. MA1's U subclade was just as related to the type Mesolithic Europeans had as north African U6 is to them. Why would two different population have the same mtDNA haplogroups? That doesn't make any sense.

I will have to read over Laz 2013 again to see if they ever mention a deep relation with WHG and ANE. Being hunter gatherers means nothing, mixing with each means nothing, being connected with Indo Europeans means nothing. Just because Spainish and native Americans mixed in colonel times doesn't mean their closely related.

Grey said...

Okay, i'm not an expert so 99.99% chance i'm wrong but...

i don't think they can say La Brana was dark unless they definitely crossed off the red hair option (modern MC1R).

This is how i think it works
1) early skin lightening genes (minor effect) (except *maybe* the red hair one)
2) late (agriculture time) skin lightening genes (large effect)
3) crossover effects from the independently evolved East Asian lightening genes effecting Euro hair and eye color

so La Brana has some of the earlier ones but does't have the late (agriculture) large effect genes so should be dark - and personally that's what i thought before anyway as i think the mammoth steppe population were likely to look like less East Asian versions of Native Americans


i think the red hair gene is a kind of quick and dirty skin lightening gene like a kind of red-haired albinism and i think the effect on euros who didn't have the later genes might have been more dramatic than it is today because today most red-haired people will *also* have the more recent large effect genes.

So i think WHG from that time - if they're not Cheyene they're red heads.

Krefter said...

Grey, those are genes can you mention the SNP's and the alleles of those SNP's.

Figuring out hair and eye color seems pretty easy because if it is accurate in nearly 100% of European samples you know it is probably accurate in La Barna-1 and Loschbour. Both probably had a combination of blue eyes and black(maybe brown) hair even though today in Europeans black hair usually means brown eyes. The dark skin is hard to believe since even light skin today correlates with WHG ancestry.

When will people finally realize probably none of those light skin mutations are Europeans and all were brought to Europe from the near east in the Neolithic? The same mutations in near easterns obviously don't have the same effect so now we have to find out what else causes pale skin.

Maybe the near eastern light skin mutations(most) which were brought to Europe in the Neolithic only take you from south Asian black to near eastern brown. While there were mutations in Mesolithic Europeans which took you from south Asian black to Mesolithic white. 25-35% of the skin color difference between Europeans and "Africans" is caused by rs146654 A/A not a big deal.
Not many ancient writers mention red hair. In for example ancient Greece or anywhere in the near east there would be almost no reason to mention hair color because many probably thought everyone had dark hair. In the Bible they never mention hair colors except old men have grey hair and young men have black hair.

Tacitus said all the Germans have red hair(but we know that isn't true) many ancient Greeks said the Thracians had all red hair(but we know that isn't true) Herodotus said a Scythian tribe in Ukraine the Budni had all red hair(but we know that isn't true), Ibn Falden said all of the Rus Vikings had red hair(but we know that isn't true). An ancient Chinese writer said all of the Franks have red hair(but we know that isn't true). The Romans mentioned red hair in Picts more oftenly than any hair color but it was probably only 10-15%.

The reason red hair was mentioned in people who had it is because it sticks out to people who have never seen it before or rarely ever do. I doubt in most physical descriptions of those people that they ever mention red hair just you hear about the ones that they did.

I think it is likely that red hair in west Europe is connected with Y DNA R1b L11 and its presence in Indo Iranians-Tocharian's and its peak in Udmurts could mean it spread out of Russia with some Indo European languages.

Grey said...


"Tacitus said all the Germans have red hair
many ancient Greeks said the Thracians had all red hairHerodotus said a Scythian tribe in Ukraine the Budni had all red hairIbn Falden said all of the Rus Vikings had red hair
An ancient Chinese writer said all of the Franks have red hair
The Romans mentioned red hair in Picts more often"

Yes they did. All of them. Independently.


"The reason red hair was mentioned in people who had it is because it sticks out "

Or because it *was* true but isn't now.

Krefter said...

Can you afford to read the paper on la Brana-1?

If any can please post as much info as you can? Actually you should post the entire paper.

Krefter said...

There is a very low chance that 2,500 years ago red hair was significantly more popular than it is today. Germans were not Mesolithic Europeans none of them were. There is really low chance red hair even reached 1% in any part of Mesolithic Europe(except maybe eastern).

Grey said...

After reading up on skin color genes i think there is an extremely high chance that red hair / pale skin was a quick and dirty skin lightening fix which has gradually been replaced across Europe by later versions that allow tanning and hence mainly survives in those parts of Europe furthest from the source of the replacement genes.

Grey said...

ugh, dunno, reading academic papers makes my head hurt. i prefer them pre-digested by bloggers. if i can't find anything about if La Brana had the red hair gene i'll think about buying it.

Grey said...

Ah found.

"SLC45A2 and SLC24A5 are classified as having the strongest colour-draining effects in Europeans
MC1R, KITLG, IRF4 with medium effects
ASIP, OCA2, TYR and TYRP1 with weak effects.

The only derived alleles found in La Braña 1 are present in IRF4 and two of the weak-effect genes, TYRP1 and ASIP."

So apparently not the derived MC1R that causes red hair - back to Cheyenne then.

Grey said...


"IRF4 is a transcription factor that has been implicated in acute leukemia.[5] This gene is strongly associated with pigmentation: sensitivity of skin to sun exposure, freckles, blue eyes, and brown hair color."


I'm back towards Cheyenne looking again but in the process some interesting thoughts: dark or light or both at once?

Anonymous said...

Pyromatic: "Why? What is your age estimate? What method do you use to arrive at it?"

I don't believe in age estimates. they are guesses based on flimsy ground and generally arguably wrong basis. They have been consitently wrong.
I followed regularly the human genetics' community for years and many were jeering, laughing and overly arrogant because of their certitudes about it.
Now, they are not jeering and laughing anymore. They learnt their lessons and just wait for aDNA.

"Are you not a fan simply because it contradicts some belief you may hold?"

Oh, come on! It sounds like you are the one that have a problem with another time frame for this hg.

Fanty said...

There is a different way to see it:
If you watch PCA or MDS then you can see that it picks up WHG against EEF as maximum avaible allele contrast (1 dimension). It does that even if there are only modern people. Its responsable for the North-South contrast.

ANE is involved in the second dimension (west-east contrast).

Krefter said...

Loschbour had the derived rs12203592 T/T(in gene IRF4) and according to 8-plex eye color predictor if someone has rs1291382 G/G and rs12203592 T/T they have blue eyes and Loschbour had both. They also listed 5 mutations and if someone has two of them they have light or medium skin. The only one Loschbour had was rs1291382 G/G that same mutation was listed in the must have mutations for light skin.

So mutations associated with blue eyes I guess are also associated with pale skin and for rs12203592 T/T(in gene IRF4) sensitivity of skin to sun exposure, freckles, blue eyes, and brown hair color.

I don't know how all these mutations work together and cant make any conclusions based on them. Obviously today light eyes-skin-hair are connected with Mesolithic ancestry and skull shape. It is still hard to believe Loschbour and La Brana-1 had dark skin.

The idea there were rapid selective sweeps that created the pale skin we see in Europe today sounds crazy to me. Why is everyone in Europe basically pale skinned? When many haven't been very connected genetically since the Neolithic? For example Portuguese don't share much common ancestry with Finnish besides Neolithic farmer ancestry and maybe some common copper age ancestry from eastern Europe. How do you explain the connection with light hair-eye-skin color and WHG skull shape? Does it just happen that all Europeans with those features have eastern WHG ancestry? How do you explain the connection between light hair, light skin, and light eyes in Europe today? Why would it be light eyes, dark hair, and dark skin back in the Mesolithic?

There are many errors in the hypothesis that light skin and hair did not become popular till the Neolithic. If it is true that light skin and hair became dominate during the Neolithic how do you explain the things I named above?

Grey said...

"They also listed 5 mutations and if someone has two of them they have light or medium skin. The only one Loschbour had was rs1291382 G/G that same mutation was listed in the must have mutations for light skin."

Yes, but they're looking at modern populations. What were people like when they had the light eyes and freckle genes but not the later skin lightening genes from the near east?

Regardless of La Brana i wonder if the earliest European depigmentation happened via the "light eyes and freckles" quasi albino genes.


"The idea there were rapid selective sweeps that created the pale skin we see in Europe today sounds crazy to me."

I think it happened twice - the quick and dirty quasi-albino way and the later farming-spawned way.

It doesn't look like the farming one is predominantly about skin color. Skin color is a side-effect. I think the second one is about breast-milk and iodine but that's a separate argument.

After my reading i'm 50/50 again: either Cheyenne dark but with blue eyes as a fluke or heavily freckled and blue eyes in a package from partial depigmentation.

Unknown said...

I don't understand how someone can totally disregard the statements of a professional in the scientific community. A paper that they haven't read to boot.

If dark skin and blue eyes were in Europe, so be it. If light skin came with farming, not a problem. Don't be surprised if blonde and red hair come from the east.

Don't be hasty to deny what people with advanced degrees, who are trusted to do this, have to say. Let it be and move on.

Krefter said...

"Yes, but they're looking at modern populations. What were people like when they had the light eyes and freckle genes but not the later skin lightening genes from the near east?"

I know that's a good point. I don't think we can judge how certain mutations made La Brana-1 and Loschbour look but we can at least guess.

All types of light pigmentation in Europe are connected with each other no one would argue against that. Why would they not have also been connected with each other in the Mesolithic? I don't understand why light eyes would go from usually being in dark haired and skinned people to being in light haired and skinned people.

The selective sweeps seem very unlikely to me. Still, the one answer I can think of that can explain why Mesolithic ancestry correlates with light hair-eyes-skin is because the most light pigmented and very Mesolithic descended people who live in north and east Europe got almost all of their Mesolithic ancestry from eastern Europe via Indo European and Uralic migrations in the metal ages.

Even Davidski considers this idea and you know he is probably very conservative about making big hypothesizes without strong evidence in his articles. After saying light skin and light eyes(I think he meant hair) probably are recent in Europe and formed after the hunter gatherers and farmers mixed and their hybrid DNA went through selective sweeps.

He said this
"But perhaps this isn't the full story, and present-day European pigmentation traits are also sourced from a late migration into Europe of a prevailingly blond people from somewhere in what is now Russia?"

I doubt the light hair-eyes-skin is connected with ANE(pretty even distribution in Europe) but instead is connected with east European WHG. I don't know of any studies but I doubt Sardinians pigmentation is much different from other southern Europeans and Sardinians are nearly full blooded Neolithic west Europeans with no east Europeans WHG.

I doubt Mesolithic east Europeans had the same light skin mutations as Neolithic Europeans, modern Europeans, and modern near easterns. But I do think they had pale skin from another source. There is skin color difference between modern north Europeans and near easterns how do you explain that? Maybe someday the mutations behind the very pale skin in Europe will be discovered.

Krefter said...

All other evidence says it makes no sense for a people to be blue eyed and dark skinned. I shouldn't just say well alright and move on. I have to find out how this is even possible and if it is actually true. It is good to be a critical thinker and to consider all the possibilities. Of course I consider the dark skin thing but since that is the main stream believe I am looking at the evidence it isn't true. I am trying to look at all the different possibilities.

The professional's have to have sources from what other professionals say. That is why Laz 2013 said so little about Y DNA. They are very conservative about what they say. From what I have heard the paper on La Brana-1 is basing his dark skin on the fact he didn't have the mutation associated with European light skin. They just went along with what has been said about these light skin mutations.

I never said they are wrong just that there are other possibilities. I see it has very unlikely dark skin-blue eye combination was ever popular.

jackson_montgomery_devoni said...

It is starting to look like the common shared ancestry among indigenous European Upper Paleolithic and Mesolithic hunter-gatherers and Upper Paleolithic Siberian sample MA-1 may be mtDNA haplogroup U. The Y-DNA seems to be more diverse but they when it comes to mtDNA they are all members of haplogroup U.

Grey said...


I'm not disregarding what they say. They may well be right. However in the course of looking at this myself i started to wonder a completely separate thing about the history of skin lightening in Europe i.e. whether there was an early version (MC1R / IRF4) based on freckling rather than shading and a second later near eastern one based on shading and going by the testimony of a long list of ancient writers i think wondering that makes perfect sense.

Whether that is relevant to La Brana or not is a separate issue.

Grey said...

"I don't understand why light eyes would go from usually being in dark haired and skinned people to being in light haired and skinned people."

IRF4 is associated with dark hair, light eyes and *freckles*

that's what made me wonder - maybe it's like an earlier / different version of the red hair "type" i.e. pale and brown skin (freckles) both at once rather than shaded skin.

I just wonder if having half white skin (i.e. freckles) might be a compromise between getting more UV but not too much?

Fanty said...

What do you think that Germans had been if "none of them" was related to Mesolithic Europeans?

What we know is, they must have been totaly different to anything that lives in the mediteranean. Tacitus claims, the Germans are like no other people in the known world. They dont resemble anyone, but themselfs.

And the Romans knew a lot of people. They even knew the Finns. And Tacitus says, some think the Finns are Germans and others think they are Sarmatians (Russians), but he thinks they are none of the two but a people on their own. Remarkable for a people who thought Scandinavia is an ISLAND in the "Mare Germanicum" settled by a seafaring version of Germans.

Chad said...

Just because something doesn't fit how you want to see the world, doesn't mean it doesn't exist. I have friends who are half black, and have blue eyes. They are just as dark, if not darker than La Brana.

Blonde hair, blue eyes and light skin did not start together. Mesolithic Europeans and Mal'ta do not have light skin alleles. This also does not mean that all Europeans had blue eyes then. There's no need for extremes.

The genes linked with blue eyes are only minor in skin color. La Brana had one or two small ones correct? Not enough to change his skin from dark to light, sorry. You may not notice the difference from the genes, depending on his level of tanning.

As for the ridiculous comment about there not being any blue eyed Native Americans, there can't be dark skinned blue eyed people is ridiculous. Blue eyes almost certainly did not exist before Native Americans had already arrived in North America.

Also, there is no reason for people to have light skin, even in a dark environment as long as their diet is rich in vitamin D. Look at Siberia and North American natives with darker hair, eyes and skin. I see no reason why Mesolithic Europeans can't be the same or slightly darker than Native Americans. It's not coal black, for god's sake!!

Grey said...

" La Brana had one or two small ones correct?"

La Brana had IRF4

"This gene is strongly associated with pigmentation: sensitivity of skin to sun exposure, freckles, blue eyes, and brown hair color."

It might be nothing but that's what made me wonder about the freckling option.

Krefter said...

Chad, I am fine with Mesolithic Europeans being full blooded Nigerians. I just think it is bad to assume La Brana-1 and Loschbour had dark skin it does make sense today.]

Sure you can give examples of rare individuals today with dark skin and blue eyes but you cant give any evidence that it is common. Look at the maps even light skin is connected with light hair and eyes. Have you done any research or do you just assume. Over 70% of Germans with black hair have brown eyes and over 70% with blonde hair have light eyes. Stop ignoring all other evidence.

Chad said...

freckling is not a big deal. some African Americans have freckles. A little bit more reasoning why light skin is not needed pre-Neolithic for you.....

Vitamin D is needed to digest calcium.. when did Northern Europeans start drinking lots of milk? Who is the most lactose tolerant? Northern Europeans, correct? Some pressure was put on Europeans to be able to digest milk, pushing skin lighter to bring in more vitamin D.

Perhaps all of the milk drinker's are the ones that survived some type of viral event due to animal domestication. The selective pressure would've been pretty high. Post heavy meat and liver diets would need a new source of vitamin D. Light skin is key to absorbing Vitamin D and in turn digesting calcium. Perhaps the more lactose tolerant we got, the lighter our skin got.

Chad said...

Or visa versa on the last sentence. I don't care about current phenotypes. We did not look like that 8kya, or especially 24kya. We don't even know when or where blonde hair began, so don't get the cart before the horse.

Chad said...

I believe the oldest blonde specimen to date is from the steppes about 3.2kya, I believe. Davidski could probably highlight this better. This steppe population had dark eyes, darker skin than current Europeans and Light or blonde hair.

Krefter said...

Chad, how do you explain such similar distribution of light hair-eyes-skin in Europe to Mesolithic ancestry? How do you explain that light hair almost always means light eyes and light skin?. You get what I am saying.

first you have to study the population history of northern Europeans don't assume Funnel beaker people are Scandinavians ancestors because now we know they were not. Almost all northern Europeans ancestors arrived where they are now within the last 6,000 years. They were probably already as pale as they are now. Evidence of that is the Andronovo people and other bronze and iron age Indo Iranians who had vast majority blonde hair and light eyes. I guarantee you plenty of blondes will be found in Corded ware culture dating back to 3,000BC. You have a time frame from about 3,000BC-2,000BC for every little north European population to developed light skin. Do you see how unlikely that is?

Did they have dark skin, blonde hair, and light eyes before 3,000BC? Your theories defy logic. I am not saying that whole thing with vitamin D is not a good idea but when looking at all other evidence it seems impossible.

Give me one piece of evidence besides La Brana-1 and Loschbour that light skin, light hair, and light eyes are not connected to each other and Mesolithic European ancestry? Can you name one modern population that has a high amount of blue eyes but also dark skin?

Grey said...


"freckling is not a big deal."

Not usually except *maybe* in one situation where it might have been a very big deal.


"Light skin is key to absorbing Vitamin D"

Yes and there is a trade-off - you need light skin to get more UV but you mustn't get too much. So what is the best trade-off in that situation?

(before tanning)

Is it a halfway shade between light and dark skin or is it half light skin and half dark skin i.e. freckles - the white skin to get more UV but the dark freckles to not get too much?

I don't know but it's an interesting idea.

Chad said...

Sorry for another post folks, but all it takes is one blue-eyed person to put the genes in a population. It's a mutation, not a recessive trait.

Perhaps the blue eyes took them from being even darker than La Brana, down to where he was. Skin doesn't just go from dark to light. It might've taken Oetzi's ancestors all of that 5000 years of farming up to him to get the olive skin.

Krefter said...

Loschbour had no MA1 related ancestry. I don't think it is a direct relation but the only relation there is comes from mixing between the two.

mtDNA U doesn't=hunter gatherer. The farmers brought some U lineages such as K and U3. There is only one mtDNA sample of ANE people and it was U-MA1(MA1 was apart of his own subclade that today is extinct).

Krefter said...

Chad it seems you never consider genetic when talking about pigmentation history. Think about the amount of common ancestry all modern European groups have. 5,000 years after arriving that is more than unlikely it is impossible.

Dark skin blue eyes in hunter gatherers light skin brown eyes in farmers that MAKES NO SENSE. Consider how WHG ancestry is connected with light pigmentation in Europe while EEF ancestry is the opposite. Consider how light hair almost always means light eyes and the other way around and how both are connected with pale skin. Why do you ignore these things?

Krefter said...

Grey could you list La Brana-1's pigmentation genes? That would help alot

Grey said...

quoting from upthread

"SLC45A2 and SLC24A5 are classified as having the strongest colour-draining effects in Europeans
MC1R, KITLG, IRF4 with medium effects
ASIP, OCA2, TYR and TYRP1 with weak effects.

The only derived alleles found in La Braña 1 are present in IRF4 and two of the weak-effect genes, TYRP1 and ASIP."

IRF4 is the dark hair, blue eyes, freckles gene.

(Not saying that proves anything but it's interesting.)

Chad said...

Contact the authors, explain to them that a kid without a biology credit knows more about genetics than they do.

About Time said...

@Barak "Give me one piece of evidence..."

Here is evidence:

Mal'ta 26kya Siberia = dark hair/eyes/skin + freckling

Loschbour Mesolithic W Europe = dark hair/skin + light eyes

Stuttgart Neolithic W Europe = dark hair/eyes + light skin

Catacombs Bronze Age Russia = dark hair/eyes + light skin

Kurgans late Neolithic Russia = dark hair (skin/eyes unknown)

Andronovo Siberia Bronze Age = light hair/skin/eyes

Modern = all over place in various mixes in trihybrid European pops.

We can't make any inferences about Neolithic / Mesolithic / later (!) origins of light hair. Except that it is in Siberia in Bronze Age.

Barak has theory of light hair/skin/eyes in Mesolithic Europe, but data points do not confirm (nor really contradict bc too few)

Davidski has theory that Kurgans were bearers of light hair/skin/eyes as package. but we still don't know if this was ANE (somehow changed since Mal'ta) or EEF or later origin.

Grey says light hair could be effect of ENA influx.

I think blond hair might be from a northern branch of EEF (my money is on Funnelbeakers).

Until we get more data, it's a lot of guesswork. Gok4 could quickly disprove my theory though, if I add stipulation that Funnelbeakers were mostly blond or almost uniform in this trait.

Chad said...

The European type of blonde is spread through Asia as well. It links up pretty well with IE spread. If it was Funnelbeaker, then those people would have to have their genes selectively spread to Asia. I would bet on a steppe origin somewhere between Ukraine and Mongolia. Blondism is recessive, it must be selectively bred with to spread. It is not like blue eyes.

Krefter said...

The rs12203592 T/T(in gene IRF4) combined with rs1291382 G/G(in gene OCA2/HERC2) means for sure blue eyes according to 8-plex system and it seems both la Brana-1 and Loschbour had them. It is behind shocking how similar the two are. I knew Loschbour wasn't a fluke and la Brana-1 would probably have the same results in pigmentation.

The one gene KITLG is dominate in all non Africans so probably little to no effect even though it was estimated to 15-20% of the melini difference between Europeans and Africans. Reminds me of how the form in gene SLC24A5 was estimated to be the source of 25-40% of skin color between Europeans and west Africans. Both probably mean very little. Do you see how it is so Eurocentric these studies are. They ignore the fact that the one in SLC24A5 is dominate all over west Eurasia in many people who are brownish skinned.

I am wondering if both La Brana-1 and Loschbour did not have blue eyes just we assume they did based on how those mutations effect modern people. But the accuracy of the 8-plex test makes it seem they did have blue eyes(for Loschbour blue or green).

Krefter said...

MA1 was 24,000 years old not 26,000. Your only evidence is ancient DNA.

In modern people light hair eyes and skin are ALWAYS connected. You will sometimes have rare occasions were dark skin and blue eyes mix.

HOW MANY TIMES DO I HAVE TO SAY THIS. Light hair-skin-eyes correlate very closely with Mesolithic European ancestry. Light hair eyes and skin are always connected. Neolithic European ancestry correlates with dark hair, dark eyes, and tanner skin.

Scandinavians and Baltics have the highest amount of Mesolithic ancestry and they also have majority light hair and eyes and the palest skin in Europe. Sardinians have the highest amount of Stuttgart/Gok4/Otzi like ancestry they have the lowest amount of light hair and eyes in Europe and darker skin than Scandinavians and Baltics. HOW in the world do you explain that?

You know that probably over 90% of modern Scandinavian Y DNA is descended from people who came after the Neolithic? If the farmers rarely mixed all the way up to Sweden why would they suddenly begin to mix when they get there?

I cant wait to see more detailed results of the Funnel beaker farmers and Pitted ware hunter gatherers. My bet is that the Pitted ware hunter gatherer will be very Loschbour like plus some ANE admixture and EEF. I think their Y DNA will be almost entirely under hg I and probably some form of I2a1 maybe descended of the pre-I2a1 or close relative to modern I2a1b that was in Mesolithic Sweden MAYBE I1A2 L22. They will probably also have some Y DNA G2a, E1b1b V13, and maybe some J1 and J2 from farmer admixture. I also think they will have nearly all dark hair, a lot of light eyes, and missing the so called light skin mutations just like Loschbour and La Brana-1.

For the Funnel beaker farmers I predict very similar results to Stuttgart just some extra WHG maybe like modern southwest Europeans(x Sardinians). I except mainly Y DNA G2a like other Neolithic Europeans and also some E1b1b V13, maybe J1 and J2 types. There will also probably be some hunter gatherer I2a1 or I1 types.

There was also autosomal DNA taken from an ~8,600 year old hunter gatherer from an island near Gotland named St, Forvar. Wont be surprised if he or she gets nearly identical results to the Motola hunter gatherers in near by Sweden and from about 600 years later. I wonder what pigmentation the Motola hunter gatherers had.

Grey said...

@About Time
I think they are assuming the mesolithic samples are dark-skinned because they don't have the later skin-lightening genes. However if only 1-2% have IRF4/MC1R and 98% have one or more of SLC24A5/SLC24A2/KITLG/OCA2 then the number of people who have one of the first but *none* of the second may be very small but i'd guess a few must(?) exist in places like Scotland or Ireland? If they did and they were brown and blue-eyed i think someone would have noticed but if they were pale but very freckled no-one would notice.


In theory it would be easy to check if La Brana was dark or freckled.

If there is someone in Ireland/Scotland who has


but doesn't have any of


then we could see if they were dark-skinned or freckled.

Chad said...

blonde hair is recessive, it must become a major component by sexual selection. If there was an aversion to it, it wouldn't exist today

Chad said...

Blonde hair and blue eyes do not go together. They are different genes that do not depend on each other. Blue eyes is a mutation. Blonde hair is a recessive trait. meaning it must be sexually selected to become numerous in a population. They don't just become blonde because they have blue eyes or a certain skin tone. That is ridiculous pseudoscience. Learn the difference between how blue eyes and blonde hair is spread. They are independent.

The most widely accepted reason for skin color, I have covered already. It has to do with diet. EEF is the proof. Needing to be lighter on top of that for even more vitamin D to digest calcium is another.

About Time said...

@Barak, I've given these examples before.

Irish have uniformly pasty skin that tans little if at all + Dark or medium hair (little true blondism) + lots of blue eyes. Black hair is not uncommon in Ireland and cannot be associated with recent mixture (bc nobody went there but some British mostly).

Before genetics, it was commonly thought Irish must have some kind of Mediterranean "Spanish" genes for dark hair. Well, no. They are about as northern as the English, Scottish, S Scandinavians, and Germans. Moderate EEF (not elevated).

Scandinavians have similar EEF ratio as Irish/Scottish, but lots more blondism and they usually can tan well.

Hungarians/Austrians/Germans have higher EEF than Irish, but with lots more blondism and generally can tan well also.

High EEF Ashkenazi Jewish have lots of blondism (including frequent "Scandinavian" type of blondism) and light eyes. Very little WHG and just a tad ANE.

Just some examples. It's been thousands of years, so lots of time for local sexual selection etc based of aesthetics and random local change.

We need Neolithic or earlier genotypes to date these things. But modern examples are all over the place. Modern Euros are mixed, but what they all have in common is EEF. That's where I bet blondism comes from (an early northern subgroup like Funnelbeaker).

Chad said...

Blond wasn't favored by Irish or Brits, so it is not strong there at all, they are way more dark haired, even though they have the same reflective quality as Scandinavians. Sexual selection NOT correlated!!

Davidski said...

The problem is that we don't know who those dark eyed steppe people were. The latest info we have on that, described at the link below, is very vague.

They might have been mostly individuals from the far western steppe and from the Balkans, where EEF ancestry has surely been at very high levels since the Neolithic.

On the other hand, we know that the light haired and light eyed nomads from Siberian Kurgans were closely related to present-day Eastern European Indo-Europeans because they carried Y-DNA R1a1a and Eastern European-like mtDNA lineages. They probably also had high levels of ANE because of the very high frequency of Y-DNA R among them.

Fanty said...

In a statistic from Germany, Blond hair and blue eyes come in pairs, at least in Germany.

The odds turn from blue to brown with medium brown hair.

I posted the percentages in an other tread here.

I repost it here:

Light blond haired Germans have:
71.2% blue eyes
12.1% mixed colored eyes
9.1% green
4.5% brown
3% grey

Middle blond haired Germans have:
51% blue
17.5% mixed
15.1% green
9.6% grey
6.8% brown

Dark blond haired Germans have:
42% blue
22.6 mixed
15.7% brown
12.9% green
6.2% grey

Light brown haired Germans have:
31.3% blue
24.7% brown
19.3% green
16% mixed
8.7% grey

Middle brown haired Germans have:
32.4% brown
25.5% blue
20.7% mixed
16.2% green
5.1% grey

Dark brown haired Germans have:
43.9% brown
22% mixed
16.9% green
14.6% blue
2.6% grey

Black haired Germans have:
63.6% brown
12.6% blue
11.9% mixed
9.3% green
2.6% grey

Fanty said...

If those things would be totaly independant, why does it stick together like that, after thousands of years mixing?

Davidski said...

Yep, and we see exactly the same thing among the European-like nomads of Siberia from the Kurgan burials.

So where did it come from? The same source, or are we looking at parallel selection in different parts of Europe, and even Eurasia, during and after the Neolithic? I think it's a bit of both; the same basic mix of hunter-gatherer and farmer genes being put under the same pressure by a new Neolithic diet all over Europe, followed by massive migrations during the early Indo-European period from a source more affected by this process than other parts of Europe. That's not to say that the blondest parts of Europe today are the most Indo-European genetically, because under my model it was a two pronged process, so selection might have been more important in some areas, while Indo-European ancestry in others.

Fanty said...

"If the farmers rarely mixed all the way up to Sweden why would they suddenly begin to mix when they get there?"

Well.... Farmer Y-DNA is almost limited to southern Europe while Farmer mtDNA dominates very part of Europe.

MODERN sexual taste is, that Farmers had the sexy looks. WHG looked like bouncers. Its virtually like that today... if someone is sexual attractive, then he usualy has a phenotype that the farmers introduced. And if someone has just average or even unsexy looks, he does more match WHG looks. So more unsexy, so more original WHG like does someone look. Haha. ;)

So, why would Farmers mix with Hunters? Because Hunters killed all the men and took their attractive females for themselfs? ;-P

Unknown said...

Let's not use Germany. Remember the pressure but on the populace to be these blonde and blue gods? This may skew the results. Once again, blonde is recessive. It must be bred in. Groups don't just turn blonde.

Davidski said...

There is certainly a skewed ratio of farmer Y-DNA and mtDNA in Europe today, and I think this is linked to the collapse of Neolithic societies during the late Neolithic in much of Europe, followed by the expansions of hunter-gatherer-derived groups from the peripheries of Europe to fill this vacuum.

Women have always survived such situations better, because they were basically seen as a commodity, while males were viewed as competition, and mostly either excluded from society or killed.

About Time said...

A scenario I imagine is that there were little Neolithic settlements here and there, with some tendency towards inbreeding due to isolation and need to preserve wealth/skills etc (like early Jamestown or other colonial settlements).

Some families or towns might have (by chance) had higher than usual concentrations of genetic problems, like albinism/blondism. A few bad harvests, reduced health all around, illness/plague. The "oddball albino" families were healthier due to Vitamin D advantage and survived when others didn't.

Word spread, and suddenly they became more popular. The "ugly ducklings" became desirable mates, maybe viewed with a slight sense of awe because they were "special" to survive a devastating plague. Just a scenario.

About Time said...

@Fanty. If that's true about modern taste (if - I have no opinion personally), it's maybe to do with facial features that show emotion and attention better.

It's also notions of classical beauty and a sculpted bone structure that shows each feature in a harmonious way. In other words, in terms of perception (i.e., how the brain looks at a face), a cognitive/sexual preference for clearly delineated features.

In other words, I think of it as sexual/social relating to the brain and social cues (which are, bar none, the most important human adaptations due to ability to cooperate, according to Darwin) more than intrinsic properties of skeletal anatomy.

Grey said...

"Blonde hair and blue eyes do not go together."

OCA2 influnces blue eyes
KITLG influences blond hair

The genes are different but if the genes tend to go together for a separate reason e.g. they came originally from the same place and were brought to Europe by a particular population then their distribution might correlate.

I think that's what happened.

(I think they're likely to be originally East Asian btw.)


"Blond wasn't favored by Irish or Brits, so it is not strong there at all"

Exactly, pale and red with freckles or pale and dark with freckles.

That's why i think skin lightening in Europe happened twice: an early quick way with freckling and IFR4/MC1R and then the later near eastern way with SLC24A5 etc.

(Three times if you include Neanderthal.)

Grey said...

Say for the sake of argument the original coloring was

1) Original WHG/ANE: dark hair, dark eyes, brown skin
2) WHG (after IRF4/MC1R) was dark or red hair, light eyes, pale skin but very freckled
3) EEF: dark hair, dark eyes, light but tannable skin
4) ANE: same as WHG plus OCA2/KITLG (so much more blond/blue) and possibly plus the near eastern genes from EEF somehow

Then say
1) WHG survived most in the Northwest corner: Isles to Baltic
2) EEF expanded roughly south-east to northwest
3) IE expanded roughly north-east to southwest

Then the Dutch would be at the join of all three WHG + IE + EEF
The Baltic would have lots of WHG + IE
The Isles would have the most unmixed WHG

oldtimer said...

If it helps - the C1a2 person who has proven Irish/English heritage back to mid 1600s has blue eyes and very very fair skin, does not tan and has freckles.... there is also lactose intolerance in the family.

Fanty said...

"Once again, blonde is recessive. It must be bred in. Groups don't just turn blonde."

The same was claimed for blue eyes. That original Danish paper about how blue eyes came to existance claimed that blue eyes are recessive and could have only reached levels of 70+ in the northern half of Europe because of sexual selection.

Grey said...


Fanty said...

German taste was checked some years ago.
Under several other things averaged faces of the most attractive 4% and the least attractive 4% had been done to have "prototypes" of sexy and unsexy to see what the difference may be,

The difference was:

- sexy face is slim, unsexy is wide
- sexy is dark haired, unsexy is blond
- sexy is tanned, unsexy is pale
- sexy is low forehead, unsexy is high, big forehead.
- sexy is high cheekbones
- sexy are edgy outlines of the face, unsexy is roundish
- sexy are small but dark colored eyebrows, unsexy are big or light colored eyebrows

Thats from their text... through I cannot find some of those differences in the images...

I putted those "Prototype" images they did with the reconstruction faces and some modern guys on one image here:

Unknown said...

A modern persons phenotypes matter none.

@ grey
Blue eyes were in euro before blonde hair and red hair. They were tested for red hair like loschbour. ZERO chance of blonde or red hair in their alleles. Blonde hair likely started in the east or post Neolithic, and had to be selectively bred for. You cannot be blonde without your parents carrying the allele. Even then it isn't a guarantee. It was passed to me and I have dark hair.

@ fanty
Blue eyes are not recessive

I'd bet all my money on milk digestion bringing on pale skin we see in Northern Europe.

You need the dysfunctional mc1r to have a chance at pale skin a red hair. This gene is not involved in the evolution of pale skin. Look it up.

Unknown said...

If everyone wants to follow maps of current maps of phenotypes, you're traveling down the same crackpot road of the r1b ice age refuge people. Maps don't mean a thing without data! Zero chance of blonde or red hair in all mesolithics tested! ZERO!!!

About Time said...

@Fanty: I think the "sexy" features relate to face recognition. Not kin recognition or individual recognition, but features key in recognizing a face as a human face. And for showing emotions and thoughts.

In other words, selection for a more "face-like" face. With fewer obscuring features (heavy brows, rough lines, etc) and each feature clearly visible (eyes, nose, mouth, eyebrows, etc).

Almost like if you had to optimize a face type to be recognized by a computer with clearly legible expressions.

Exaggerating these features (runawayy selection in Darwinian terms) could also happen at expense of beauty sometimes (which is a luxury in nature anyway). Still social effect - the main benefit - is preserved.

I think this type of selection happened in Basals at the same time other non Africans were selecting to become more efficient hunters. Hunters did not select so much for this. Projecting too much feeling and thought can be disadvantage in many situations. Hunters might have preferred a harder "game face" or "warrior face."

Darwin said man is a weak animal whose main adaptation is in group cooperation amd mutual care. Basals (in my scenario) optimized aspects of this quintessentially human capacity. Others later found it compelling and attractive,

Davidski said...

Here's my favorite hunter-gatherer-like celebrity. Interestingly, her eyes are mixed, but the face is definitely hunter-gatherer-like.

She's way more attractive than this model IMO, who probably has more Neolithic phenotype alleles (but at the same time, light eyes).

Here's what the Copper Age steppe people looked like IMO.

Now that's what I call an analysis. :)

Krefter said...

Davidski, I understand what your saying and this is my response.

All theories of powerful selection to create modern European "coloring" make no sense to me for the following reasons.

>Light hair-eyes-skin are connected to each other. EX: Blonde hair almost always means light eyes and light skin and Black hair almost always means brown eyes. How could it have been the opposite in the Mesolithic and switch in all of Europe.

>Mesolithic ancestry correlates with light hair-eyes-skin, while Neolithic or any other form of near eastern ancestry is the opposite.

>Not all Europeans have significant amounts of common ancestry since after the early Neolithic. For example Sardinians and Finnish.

>There is a good chance Stuttgart and all other early European farmers had the same skin tone as modern southern Europeans. I guarantee you blonde hair anywhere is southern Europe almost always means blue eyes, and brown skin almost always means brown eyes. Even for Sardinians who probably have some Loschbour related ancestry. Obviously blonde hair was not selected in them but I guess you can say light skin was selected separately all over Europe.

>I don't understand how it is possible a mix between Loschbour like people and Stuttgart like people+selection=Baltic and Scandinavian pigmentation. We know there were people pigmented like that at least by 3,000BC.

I think the most probably solution is extreme paleness represented best around the Baltic sea and in ancient Indo Iranians is from Mesolithic east Europeans who's blood spread via Indo European and Uralic migrations during the metal ages.

Grey said...


IRF4 is like MC1R but for **dark** hair, light eyes and freckles.

It will be provable imo. Even if IRF4 is 1% and SLC24A5 is 98% among European descent populations there might still be a few people with IRF4 but not the later skin lightening alleles (and in the past there would have been lots of people like that).

Grey said...

If the alleles that separately increase the chance of blond hair and blue eyes both came with the same population from the same direction then the frequency could correlate without being related.

Krefter said...

Grey, you make the assumption WHG and ANE are the same when they are not. Even after not mixing for over 24,000 years you think both developed the same mutations for light eyes?

About Time I do know that WHG distribution does not perfectly match the distribution of light hair and eyes and that dark haired and brown eyed or light haired and light eyed percentages can vary.

It is true that Irish are most known for having very black hair or red hair, blue or green eyes, and pale white skin. At times I can easily recognize Irish Americans.

According to this

Only 38% of Irish have light hair(including 9% red) and 82% have light eyes(including 66% blue). That is very different from Swedish who have 78% light hair(3% red) and 89% light eyes(72% blue). The Welsh(also Insular Celts) have also been known for very dark hair or red hair and very light eyes. According to that link 42% have light hair(11% red) and 68% light eyes(50% blue). I remember reading ancient Roman stuff about the Britons they said they were very dark haired and the Gauls were more blonde haired. They even said they look similar to Iberians. That is constant with the current pigmentation of modern Insular Celts. I am sure highlander Scots are always mainly dark haired and light eyes with about as much red hair and blonde hair.

You can tell that hair and eye color percentages have changed over time because of random sexual selection or whatever. So I guess it is possible(but very unlikely) blue eyes were once popular in dark skinned and haired people. But why is it not like that pretty much every where except for in Insular Celts? Another thing about pigmentation of Insular Celts is even the Romans had mentioned on their occasional brown skin. I have seen it before and I have seen other naturally brown skinned north-west Europeans.

Chad said...

But there is not blonde hair in the Mesolithic Europeans... so that argument does not work......

Chad said...

These are not amateurs. They understand which genes cause lightening and what the percent of change is. If there were additive genes that lead to lightening, which we see in Europe, they would know. They found it in the Neolithic farmers. It wasn't in the Mesolithic. Acceptance of this fact, will allow us to speculate on where blonde and red hair come from and not argue about what has been decided.

It is not unlikely for these Mesolithic people when admixed with farmers to have become nearly as light as farmers. Later, the additive genes found almost only in Europeans could be added on to make the changes we see today. People could've still been the color we see today by the Iron Age. Gone before any written language in Europe, therefore all memory of these dark skinned people was lost. Surely by the Iron Age and maybe the Bronze Age there would likely not be a pure Mesolithic person left.

Chad said...

Where are you seeing IRF4 affecting skin tone? Adding a chance of freckles, blue eyes and down to brown hair, does not make a pale northern European. You need several other alleles at play. Some of these that affect skin color have up to 30 difference alleles involved in the changes. You need more than just one SNP on one minor gene. Blue eyes and up to 25% chance of dark brown and not black hair. Still not pale... still not blonde....

Davidski said...

So if WHG and ANE didn't mix for 24,000 years, then how come Motala12 was classified as 19% ANE?

Grey said...


I'm not saying there was. I'm saying if both alleles came into Europe among the same population from the same direction they could correlate in later populations without being directly related. For example if they came in from the northeast with the IE then they could both share the same cline.

Example of what i mean with two separate alleles but with the same frequencies along a cline

Northeast: 60% frequency each -> 24% chance of both
Central: 40% frequency each -> 16% chance of both
Southwest: 20% frequency each -> 4% chance of both

Unknown said...

Yes that is possible, but you still get sexual selection to play with for 7000 years.

Mesolithic Europeans were likely somewhere around australoids and Papuans in skin color, with dark brown to black hair, which I believe is the same mc1r change as amerinds and then blue eye mutation.

The question is what did the climate in the north, milk, and grains do to make such a change. I bet sexual selection played into skin as well. Maybe 4000 years ago we picked up a gene to like blonde and blue white girls. I've read stranger stuff!

Grey said...


"Grey, you make the assumption WHG and ANE are the same when they are not."

**Were** the same (or very closely related) at one point. I think WHG **were** the southern forest dudes during the LGM and the ANE **were** the mammoth steppe dudes and they are related somehow.


"Even after not mixing for over 24,000 years you think both developed the same mutations for light eyes?"

Apparently there are many versions of the MC1R mutation that creates red hair and light eyes and freckled skin. That implies the same mutation has happened multiple times among different populations. I think it's stuck around among muliple northern latitude populations because it is a quick and easy way of increasing UV.

(The Jomon Japanese have it too - convergent evolution.)

To me it looks like the IRF4 mutation does the same as MC1R but with dark hair instead of red.

Unknown said...

Blue eyes won't be related to the blonde and pale. I'm speaking in regards to their appearance. Blue eyes developed separately. Mesos are proof of that. Blonde and light could come in together, but by then most everyone should have EEF doing some lightening. All instances of these being together are sexual selection and maybe some climate and diet issues.

spagetiMeatball said...

I don't see why you think the second woman looks neolithic.
Here's what I think the farmers looked like:

just without gold teeth. This is the teeth?

And this is what the WHG's looked like:

First one is some turkish farmer, second one is the guy who played William Wallace's buddy in braveheart. He looks pretty similar to the reconstruction of the Brana guy in this paper.

Grey said...


"They understand which genes cause lightening and what the percent of change is."

Do they? If IRF4 is 1% and SLC24A5 is 98% then people who have one but not the other are going to be very rare and therefore easy to miss.

(Effectively you're looking at difference in base coat. Among Indians the base coat is brown and then the differences of the various alleles can be calculated on that basis. If the base coat was white with freckles i.e. half-white and half-brown then the effect of the other alleles might just be to paint over the freckles.)

If the freckle idea is correct in the past people like that would have been a lot more common, especially in places where the frequency of SLC24A5 increased the latest e.g. the Isles. And yet no record of lots of brown people with blue eyes.

Anyway, like i say there should still be a few people like that living today and so we can see if they're brown or very pale with lots of freckles.

Grey said...


"Gone before any written language in Europe, therefore all memory of these dark skinned people was lost. Surely by the Iron Age and maybe the Bronze Age there would likely not be a pure Mesolithic person left."

In Europe but not everywhere. That's partly what makes me think they're wrong about La Brana. I think the first wave into Europe were Khoisan looking and the second wave were originally Indian looking but given all the later written accounts fitting a particular kind of depigmentation (red or dark hair, light eyes, pale with freckles) I think La Brana is too late to have been the original kind and the light eyes are the clue.

Grey said...


""Mesolithic Europeans were likely somewhere around australoids and Papuans in skin color, with dark brown to black hair"

Originally i think they were Indian looking (but more archaic) with dark hair, dark eyes and dark skin (like MA1?)

However I think the IRF4 and MC1R mutations (combined with ASIP and TYR or TYRP1) made them dark or red haired, **light** eyed (not necessarily blue) and pale skinned with freckles.


"Blue eyes developed separately. Mesos are proof of that."

Light eyes: blue, grey, green etc.

The written records mention red hair with green eyes and red hair with grey eyes etc.

Davidski said...

Whoaaa...I didn't say Anja was a straight up Neolithic farmer from the Near East. I just said she had more Neolithic-derived phenotype alleles than Magda, so she looked more like early European farmers than her. It's pure speculation on my part, but I think generally correct. The one I'm not sure about is Gosia as an early Kurgan steppe nomad, but she has a kick ass look so why not?

Grey said...

Actually the survival of any early de-pigmenting trait is inversely proportional to distance from the east and southeast then Norway and Iceland might be possible locations also.

A quick google of "norway" and "freckles"

and some extreme freckle faces: 50% brown, 50% white?

less than 50% but still high

and just cos hubba

Grey said...

meant to say "Actually **if** the survival..."

About Time said...

Dienekes has a new thread about a paper showing Neanderthal origins for some skin pigment genes:

"Analyses of surviving archaic lineages suggests that there were fitness costs to hybridization, admixture occurred both before and subsequent to divergence of non-African modern humans, and Neandertals were a source of adaptive variation for loci involved in skin phenotypes."

Not sure which skin phenotypes those were, but throwing that in there.

About Time said...

Mesolithic Euros might have been varying shades of brown, just like most people around the world. Thinking if Cheyenne or Indians (from India) makes sense IMO.

If Basals were Natufian, they might have picked up some darker adaptations near the equator. Coming into West Asia, ebony-mahogany Basals might have faced discrimination from the brown skinned locals (color prejudice is pretty common in Arab world and India even today). Maybe they started selecting for lighter skin and ended up overcompensating by becoming not just brown or olive, but pale tending. Like the myth of Pandu.

Looking at the map, this could have happened somewhere in a West Asia (Gedrosia). By the time the got to Europe as EEF, they would have brought in the light skin adaptation.

Unknown said...

There isn't a cline to the west. Brits and Scandinavians have the same reflective rating to their skin. I think climate/cloud cover positively selected the lightest feature. Interestingly the highest lactose tolerance is with these people. Whether or not it's related is the question.

Unknown said...

I think it was the east Eurasian light genes.

Grey said...

"Mesolithic Euros might have been varying shades of brown"

I've no doubt they were originally brown as there's no reason to shed pigment until you get to the northern latitudes. It's just a question of when the de-pigmentation began. For it to have been farmers it would need to have reached every remote mountain valley with enough time to spare for no written record.

Grey said...

There's a cline from the southeast to the northwest in the red hair / light eyes / freckled light skin (MC1R) and the dark hair / light eyes / freckled light skin (IRF4) phenotypes which i think is an earlier depigmentation method.

If you've ever spent any time in the remoter parts of NW Europe you see this phenotype a lot and sometimes you see one with a mass of freckles to the point where they are 1/2 brown.

Matt said...

The Neanderthal BNC2 variant haplotypes are reportedly at high frequency in both present day Europeans and Asians.

BNC2 has been implicated in skin pigmentation / freckling variation in at least a Dutch population and web based sampling (google if interested).

Be interesting to look at Oceanian populations. Eurasian variants at KITLG seem to show their highest frequency in Oceanian populations, which causes me scepticism as to whether they are significant in terms of effect on skin tone.

An BNC2 snp rs2153271 which is reportedly freckling linked is high frequency in West Eurasians, intermediate in East Asians and low in Oceanians and Africans, as per ALFRED -

Doesn't look like BNC2 was checked out by the La Brana paper.

spagetiMeatball said...

The last model you posted (Gosia) has a very uncanny resemblance to the one of the famous tarim basin mummies, the "beauty of Xiaohe"


Beauty of Xiaohe:

This is all folk anthropology of course, but I find it hard to shake the feeling that these two ladies are descendants of the same ancient culture.

Unknown said...

It's not a method. The dysfunctional mc1r that leads to red hair was not involved in the evolution of light skin. Look at any paper on the dysfunctional gene. Mc1r is more about fur colors, it does the same on animals.

Grey said...

Yeah it is. The mutation happening so often and spreading so much - and only in high latitudes - only makes sense if it de-pigments as well.

Krefter said...

I made an error what I meant is that the relationship between WHG and ANE is they mixed and that's it. Why didn't Laz 2013 mention a very close relationship between WHG and ANE that did not involve mixing? WHG correlates with paleness not ANE, maybe they same features but not the same mutations.

Grey, In the Neolithic Swedish farmers were hardly different from Stuttgart and Otzi. The reason modern northern Europeans have so much WHG is probably because of Indo European and Uralic migrations in the metal ages. Most is probably not from Mesolithic northern Europe but instead Mesolithic eastern Europe.

Krefter said...

That is interesting but C1a2 only represents his direct paternal lineage and he probably doesn't have much ancestry from Mesolithic west Europeans like La Brana-1 and Loschbour.

Krefter said...


How do you explain dark skin in the near east and very light skin in northern Europe even though both have around the same amount of these light skin mutations? Obviously they don't=white skin. There are other reasons why Europeans have such light skin that have not been discovered!!

I know I have said this like infinity times but it seems you still don't understand.

Light hair-eyes-skin ARE CONNECTED ALMOST EVERYWHERE IN EUROPE TODAY AND IN ANCIENT INDO IRANIANS. Loschbour and La Brana-1 are confusing results and we have to find answers. Today light hair-eyes-skin correlate with HUNTER GATHERER NOT FARMER ANCESTRY. Why aren't southern Europeans the blonde ones(I don't care about random pictures on Google) since they have the most farmer ancestry? Do you see how that doesn't make any sense?

Krefter said...

About Time, just ask someone who has studied human psychology for like 50 years. Besides attractiveness has more to do with the body than anything not the face. Someone could have pretty face but be considered ugly. There are a lot of things involved and the face is definitely not the most important.

I don't get what facial features tell besides stuff that happened in peoples society and not about living and dying. Your probably not going to have a good chance of dying if your people have really wide faces but if you have snow white very untannable skin in Australia and are stone age people you will have a hard time.

The facial features an skull shape stuff is pretty interesting though. I heard some say that the Caucasian skull shape was spread by near eastern farmers but just look at La Brana-1 and numerous pre Neolithic European facial reconstructions or skulls. Obviously it has an ultra old west Eurasian source. The oldest(17,000BP) fully modern human skull in north Africa have typical Caucasian features the oldest even had the "Jewish" nose like Charchen man( very well preserved 6'6 brown haired and red bearded 3,000 year old Indo European(likely) mummy from tarim basin).

According to a map from the 1940's and today northern especially north-eastern Europeans have skulls most similar to Mesolithic "WHG" Europeans. Not a surprise since autosomal DNA has showed Mesolithic European ancestry is distributed in a nearly identical way. I wonder what the distinct features of Mesolithic Europeans were.

Krefter said...

That could be the case if almost all in Europe today is from east European Indo Europeans and Uralics. I just don't see good evidence though that light hair is from near eastern farmers. So Mesolithic east Europeans makes more sense.

Like Davidski said he doesn't have any of the mutations associated with blonde hair but one for red hair and has reddish blonde hair. Where did the blonde hair come from? Its a mystery and there is obviously still stuff to learn about where certain pigmentation comes from. If La Brana-1 and Loschbour really did not have light skin, I think they were a deep shade of brown like native Americans or south Asians.

Chad said...

There is no reason to shed pigment in northern latitudes with lots of Vitamin D. The Siberians and Native Americans are perfect examples. Please understand this point!

The general consensus is that it would take 100 generations or less to go from very dark, like La Brana to lighter skin. Nature can positively select for those traits. that's roughly 2500 years. So if the lightening starts at 6000 years ago, by 1500 BCE, we all roughly look the same shade of Oetzi or a bit lighter if selection plays more of a role. This is still before written descriptions of people in continental or northern Europe.

What this does not include is mixing of EEF and La Brana. That can speed up the process very much, not just sexual selection, but positive selection. Life is very intelligent and knows when it needs to turn things on and off. If genes are built up for grain diets and Vitamin D is not readily available, nature will find a way. I would not be surprised if all hunter gatherers of this complection were bred out by 2500 BCE.

People with writing in the Near East by 1500BCE would likely not see much of a difference from themselves. Although they may have some extra basal or African admixture by this time, so they may be darker than Europeans by a bit at this point.

Sexual selection may have really sped this up, not to mention mother nature selecting these for mixed kids.

Simply put though, without the two major skin lightening alleles, there is no way any other gene is making Europeans light complexioned before the EEF mixing. It just is not possible to go from Papuan to Swede without the SCL pair being highly involved with a few of the other 4 known contributors. There is no good biological reason for white skin to start somewhere with people eating enough vitamin D. No white skin is needed in Siberia or the Americas.

Grain eaters of the East and West Eurasians became light and spread their genes. It's pretty cut and dry there. Why there is pasty white skin some places is the tricky part.

Krefter said...

We have two nearly identical samples both probably had black or at least brown hair. That doesn't mean blonde hair did not exist in Mesolithic Europe. Like I have said more than a million times blonde/light hair correlates very well with Mesolithic ancestry today. Blondes have a much higher percentage of light eyes than any hair color.

I see it has very unlikely that blonde hair became popular in farmers.

Chad said...


This is getting ridiculous.

Near Easterners and Europeans do not share every light skin allele. You need to do better research. Near Easterners are also part SSA. That is something you need to consider.

That being said, some near easterners are close to the same color as Europeans. Look at Bashaar al-Assad and his family. His wife looks like a regular European. His kid too. You have very strange views of the world kid. You need to get out more. You have very black and white views of the world. Pun is intended.

Chad said...

I would suggest you pay attention in advanced biology, if you ever take the class. You need to learn about positive, negative, and sexual selection just to name a few things. You have no understanding of these, whatsoever

Chad said...

If you want white people to have always been light skinned with blonde hair and blue eyes, then so be it. It doesn't change the reality one bit. I'm not discussing it further with you, until you have more than your current maps that can be ruled out by sexual selection. You still do not understand that blonde is a recessive trait that must be bred for. This is why you are called pseudobama. You make wild claims with ridiculous 'evidence'. Go put your maps with the maps of r1b and tell me more about how old Europe is just like now. I refuse to discuss this with you any further until you can bring me a Mesolithic European with the recessive genes for blonde or red hair, which you wont.

Chad said...

last comment... EEFs were more likely to have black hair and brown eyes than mesolithics. .... looks solid to me as to an explanation of the dark hair and eyes of the Mediterranean and near east... There is some blondism in both places though.. mesolithics appear to have the same mc1r as Native Americans and East Eurasians that causes the hair to lighten a bit from black to a med to dark brown. Nothing that leads to blonde. Now I'm done!!

Krefter said...


All we know is that dark hair and blue eyes were probably popular in Mesolithic west Europe and dark skin is probable but debatable. La Brana-1 and Loschbour are not prove light skin or blonde hair did not exist in the Mesolithic. It seems the near eastern farmers brought the mutations associated with European AND NEAR EASTERN skin color. Some people group all Europeans as the same thing or all ancient Europeans as the same thing. The hunter gatherers and farmers were two totally different populations.

Based on La Brana-1 and Loschbour your theories make sense. But based on all other evidence I know the answer must be much more complicated. La Brana-1 and Loschbour don't represent all of Europe at that time. We will have to wait till more genomes of ancient Europeans and near easterns. Genomes should be taken of Mesolithic far eastern European(U5a dominted) hunter gatherers. They may have been very ancestral to early Indo Europeans, full of Y DNA R1a1, and a lot of ANE and WHG ancestry. I think there is also a good chance blonde and red hair will be found but not with the light skin mutations. We will also wait and see if there are other factors in pigmentation than what Loschbour and La Brana-1 were tested on.

Davidski said...

Before the comments max out, here's a study about prediction of hair and eye colour from DNA. It might have some useful info about how various markers affect phenotype.

Chad said...

Thank you Davidski,
People don't realize how blonde southern Europeans are. The percentages are not that far apart on true blondism. It's the intermediaries getting counted that distort the numbers a bit.

Krefter said...

Thanks, Davidski

Chad, we will have to get serious studies on hair and eye color percentages in Europe to understand how it varies by every little region. I guarantee you as we learn more about this mutations associated with either red or blonde hair are also associated with light skin. Same with mutations associated with light eyes.

It does seem that Loschbour and La Brana-1 were dark skinned and blue eyed but again that makes no sense for many reasons.

Chad said...

The problem with your hypothesis is that there is no light skin or light hair alleles in Mesolithic Europeans. For it to be in their populations, one of them should show the allele, as it is recessive, it must be bred to popularity like the 40% red allele in Scotland only leading to 11% red hair. If we don't find the allele, blondism does not exist there. You cannot say that blond or red hair exist in Mesolithics, when they do not have the alleles for either.

Chad said...


I posted the question to Dienekes as well, but do you know if they tested the hair texture of La Brana. I am not seeing anything. I am wondering if they have straight to wavy hair, or wavy to possibly kinky hair.

Chad said...

I assume with the pic, they must have. I hope it is not just artist speculation. Id like to see someone rebuild the face with clay, like the Stonehenge guy.

Fanty said...

"Besides attractiveness has more to do with the body than anything not the face. Someone could have pretty face but be considered ugly. There are a lot of things involved and the face is definitely not the most important."

Well, when I was young, we used to ask each others what the important parts of the girls are and ALWAYS they said: cute face is the most important, above everything else.

Someone usualy falls in love with a face.
There are also lots of words about the face (of girls in the first place)
Rough translations from the German versions:

"All it takes is a cute face and he turns into an idiot"
"He turns himself into a slave and for what? A cute face..."

Also, if one does sex in the more traditional (missionary position) way, the only part of the girl you can see, while your doing it, is her face. ;-D

Well if you make it like this:
The face may be totaly unimportant....

But if you make it like this:

Well then...

Oh... and this quiet mesolithic looking guy who plays a captured Germanic warrior in Spartacus....good body. But I still cant imagine all the girls dreaming of him. Heehehe....

omg omg...


Chad said...

From the site Davidski provided

Greek: blond (9.2%), dark-blond/light brown (37.8%), dark brown (41.2%), brown-red/auburn (3.3%), blond-red (0.0%), red (0.0%), black (9.2%)

Polish blond: (13.7%), dark-blond/light brown (44.2%), dark brown (22.6%), brown-red/auburn (1.0%), blond-red (3.9%), red (3.8%), black (10.8%)

Irish: blond (4.7%), dark-blond/light brown (32.7%), dark brown (46.6%) brown-red/auburn (6.8%), blond-red (1.7%), red (2.9%), black (4.4%)

Chad said...

so, if blonde hair means WHG, then Greeks should be more WHG than the Irish. As I have stated it is sexual selection and not an indicator or WHG ancestry. It's a sign your ancestors like getting it on with blonde chicks. That's all.

Krefter said...

Honestly your theories denial of the lists of reasons I have given you(so many times) are like denying the laws of physics. There have to be some crazy explanations for what your claiming. You don't know how blonde hair works so no prove it did not exist ANYWHERE in Mesolithic Europe. Don't you think there would be stories still today of how blonde hair grew from almost nothing to over 60% in maybe as little as a thousand years. How people all over Europe went from deep brown to snow white in a thousand or so years? There are so many other things that you ignore too. How did the old blue eyes become so inlove with the new blonde hair, red hair, and light skin all over Europe you really think that was just random? You theories if true tell are one of the most crazy makeovers in human history.

Chad said...

Greek- blue (10.9%), intermediary (12.6%), brown (76.5%)

Polish: blue (54.0%), intermediary (15.0%), brown (31.0%)

Irish : blue (50.7%), intermediary (26.5%), brown (22.7%)

Chad said...

Look at the stats I just posted. Now zip it.

Chad said...

nothing to 60% in a thousand years? try 7000 smartass. More than possible with sexual selection. You don't have a clue how this stuff works.

Chad said...

lets not forget Greeks have way more blond than the Irish but only one fifth of the blue eyes. The two are not connected but by sexual selection. I will not say it again and I am done talking to you for today.

Krefter said...

Neolithic(or post) sexual selection is not the best explanation for the distribution of hair and eye colors in Europe. According to Peter Frost every distinct European feature is the result of very strong sexual selection towards women. Are pale women really that much more attractive to cause such rapid change in pigmentation? I really doubt that is one of the main sources.

I was very surprised by those statistics considering all of the extra(non EEF) near eastern ancestry Greeks, other Balkaners, and Italians(especially southern) have. Even if blonde hair did originate or became popular in EEF it would not make sense for blonde hair to be popular in Greeks.The red hair statistics are the most surprising. Close to 7% red hair in Poland and the existence of red hair in Greece? That paper does give a great source of hair and eye color in Europe. Hopefully none of them were Americans who claim to Irish or whatever because their great great great grandfather was but the rest of their ancestry is totally mixed. I usually claim to be Scottish even though I am under 20%.

Check these statistics and also maps made by people who studied populations all over Europe and the Meditreaen.,d.aWc&psig=AFQjCNGDFKjWEyH43grctJzKnLDRnXpEjQ&ust=1391218744075581,d.aWc&psig=AFQjCNGDFKjWEyH43grctJzKnLDRnXpEjQ&ust=1391218744075581

Here I compare WHG, ANE, and EEF to hair and eye color percentages in Europe. The link under that Fanty shows how WHG correlates with all the different forms of (connected)paleness in Europe.

You should read an earlier post in this thread which shows at least for Germans blonde hair is connected with light eyes while dark hair is connected with brown eyes. From the ancient Roman and Greek writings I have read that mention physical features they had the same stereotypes about the connection between light hair and eyes with light skin(because they are based on truth). Their descriptions of ancient people such as the Britons are surprisingly accurate with modern Irish and Britons(Welsh, Cornish, and Britanny France).

French are in-between continental Germans-Insular Celts and Iberians in WHG and in hair and eye color they are also in-between the two. The distribution between overall paleness is very connected with WHG. Yet not many Europeans probably have over 50% WHG so are the origins of these features more complicated? Saying it was the way it is today in the Mesolithic + 1,000 times paler doesn't make sense when considering La Brana-1 and Loschbour.

Fanty said...

Some hair color statistics from Germany:

Light blond:
North: 4,9%
Middle: 3,8%
South: 3,9%

Medium blond:
North: 22,4%
Middle: 19,6%
South: 16,6%

Dark Blond:
North: 27,9%
Middle: 26,2%
South: 24,7%

Light brown:
North: 9,4%
Middle: 10,7%
South: 10,2%

Medium Brown:
North: 15,9%
Middle: 18,2%
South: 19,4%

Dark Brown:
North: 11,8%
Middle: 13,7%
South: 18,1%

North: 1,6%
Middle: 2,3%
South: 1,9%

Red Blond:
North: 2,6%
Middle: 2,3%
South: 2%

Red Brown:
North: 2,5%
Middle: 2,5%
South: 1,7%

North: 1%
Middle: 0,9%
South: 1,1%


Krefter said...

We know Indo Iranians that migrated out of Russia some 5,000-4,000 years ago had as much light hair and eyes as any modern Europeans. According to you blonde hair didn't exist or was popular till the Neolithic. Corded ware people will also be shown to be overwhelmingly blonde haired and light eyed and are around 5,000 years old. There is a very good chance 6,000 years ago somewhere in Russia there were mainly blonde people. Still it gives room of a few thousand years or so at the most to go from 0% to majority.

Fanty said...

I may sum it up to:

All shades rated as "blond":
North: 55,5%
Middle: 49,6%
South: 45,2%

All shades rated as brown:
North: 37,1%
Middle: 42,6%
South: 47,7%

Fanty said...

That reminds me to Roman writing about the Cimbri tribe.
They write the children of the Cimbri have the haircolor of old men.

Wich is a rather odd description if Romans would have been used to the look of light blond children (who darken when becoming older).

I was similiar too. Almost white hair with 3 years. Still light til like 10 or so.
It became medium blond until like 20. And meanwhile (41) its dark blond.

My beard is 3 colored (haha):
medium blond on the upper lip. Red on the chin and gypsi black on the cheeks. I used to say, I have the colors of the German flag in my face. X-D
Well, no longer. All black hairs turned grey. The red and blond ones are still pigmented.

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