Early European Lineages

[Angela]

This reinforces my hypothesis that blond and red hair were both brought by the Indo-Europeans (R1a and R1b respectively), but that blue eyes were already present among Mesolithic Europeans.

I have mentioned before that likely minor presence of Y-haplogroup A1a among Palaeolithic/Mesolithic West/North Europeans, but also the relatively important presence of E-M81 since the Mesolithic at least in Iberia, and possibly also in and around France. Both would have contributed to slightly darker skin among Mesolithic Western Europeans.

It's not only *these* hunter-gatherers who are darker skinned than Oetzi...so was Mal'ta...it seems to me that the evidence so far is that lighter skin did not appear in Europe until at least the Neolithic. While today, skin pigmentation correlates very well with amounts of solar radiation, this was obviously not always true. I don't think anyone yet knows the factors that brought about depigmentation, but part of the answer may indeed be that snow and ice conditions would not favor fairer skinned people. Also, until the Neolithic took hold, much of western Europe was forest. I would suggest that diet may have played a role.

I still think that diet may have played a role as well. Lactase tolerance also is not present in any of these people, and I think is undoubtedly tied to the rise of dairy farming.

 

[epoch]

It looks like WHG (West-European Hunter Gatherer) is today the truely north-european one. Most likely the Saami would have the most of it, but I couldn't find the data for the Saami in this paper.
The North-East european peak shown in K12b and Globe 13 and others probably corresponds mostly to both, WHG and ANE.

On fennoscandia they have an interesting comparison:

http://fennoscandia.blogspot.no/2013/09/la-brana-1-and-modern-european-variation.html
http://fennoscandia.blogspot.no/2013/09/la-brana-2-and-modern-european-variation.html

La Brana 2 clusters with Swedes there.

EDIT: With Sami shown separate from Swedes, that is.

 

[epoch]

It's not only *these* hunter-gatherers who are darker skinned than Oetzi...so was Mal'ta...it seems to me that the evidence so far is that lighter skin did not appear in Europe until at least the Neolithic.

Obviously that could be the case. However, Mal'ta has almost one third non-white - or rather non-european and non-west asian - admixture.

 

[Maciamo]

This actually shows that old clads can be replaced with new "improved" (more adapted) ones rather quickly and without big difficulties. It can explain quick dominance of some R1bs and I2a-Dinaric. Pretty much in agreement what you were always suspecting.

True, but where did I1 come from if not Mesolithic Scandinavia ? I think it is still a sample bias and that I1 or pre-I1 will definitely show up in other Mesolithic Scandinavian samples.

What amazes me is the incredibly wide distribution of I2a1 in Mesolithic or Neolithic Europe. Beside Sweden and Luxembourg, I2a1 has been found in Early Neolithic France (Languedoc) and was surely present all over Southeast Europe based on its modern frequency there. Many people suggested that I2a1b only arrived in the Balkans with the Slavic invasions, but if I2a1b was already in Scandinavia, the Low Countries and southern France 8,000 years ago, there isn't any reason to believe that it couldn't have been in the Balkans back then too. That would confirm an origin of I2a1b at least 10-12,000 years ago. Obviously I2a-Din (L621) is a deeper subclade (I2a1b3), but it wouldn't take 6,000 to 10,000 years for a single mutation to develop. My own theory was that I2a-Din appeared around the Carpathians during the Neolithic (Cucuteni-Tripillian culture) and spread to the the Balkans with the Thracians and the Illyrians during the Bronze Age.

 

[Maciamo]

Yet Norway has 43% autosomal DNA connected to mesolithic hunters. This strange discrepancy between autosomal connection and Y-DNA disconnection is also visible with mtDNA, I think. Norway has far less U5, U4 or U then 45%.

http://eurogenes.blogspot.com/2013/...howComment=1387851513157#c1616632050721459316

That's because H1 and H3 are also Mesolithic, not Neolithic as many people like to claim. If you add H1 and H3 to U2, U4 and U5, you get 47.5% of Mesolithic mtDNA. On the Y-DNA side there is 31.5% of I1 and 4.5% of I2a2, so 36% in total. Of course Y-DNA can get replaced by invaders more easily without impacting the autosomal DNA very much.

 

[epoch]

That's because H1 and H3 are also Mesolithic, not Neolithic as many people like to claim. If you add H1 and H3 to U2, U4 and U5, you get 47.5% of Mesolithic mtDNA. On the Y-DNA side there is 31.5% of I1 and 4.5% of I2a2, so 36% in total. Of course Y-DNA can get replaced by invaders more easily without impacting the autosomal DNA very much.

In another thread here I already wondered about this. If you look at the map of H1 + H3 you made it looks almost like the admixture map that Davidski presents. However, the more hunter-gatherers they find the more it turns out they are really almost solely U and subclades. We can now add another Swedish example and, wholely new territory, a Luxembourg.

I once thought that we mistakenly take all mesolithic hunter-gatherers for a singular entity. Since hardly any DNA was taken from France and the Benelux I figured it was possible they had all the mysterious H1 + H3. Now we have an example from the area and lo and behold: It's an U.

Still, La Brana 2 looks in some graphics like current day Swedes.

Indo-Europeans are not North-Eurasians, but Uralics are. However, the Uralic and Indo-European languages do show remarkable similarities. While some linguists consider that a genetic connection others don't. Most seem to agree though that these similarities mean they "grew up" next door to each other. Could it be that the ANE admixture may be a signal for Indo-Europeans, but that the actual IE carried only a part ANE to Europe? I am merely speculating - I am also merely a layman - but since Belorussinas and Ukranians also carry quite a large part of the WHG admixture I could imagine that parts of West-European WHG admixture was actually brought by Indo-Europeans. Just a thought though.

 

[Idun]

In another thread here I already wondered about this. If you look at the map of H1 + H3 you made it looks almost like the admixture map that Davidski presents. However, the more hunter-gatherers they find the more it turns out they are really almost solely U and subclades. We can now add another Swedish example and, wholely new territory, a Luxembourg.

I once thought that we mistakenly take all mesolithic hunter-gatherers for a singular entity. Since hardly any DNA was taken from France and the Benelux I figured it was possible they had all the mysterious H1 + H3. Now we have an example from the area and lo and behold: It's an U.

Still, La Brana 2 looks in some graphics like current day Swedes.

Indo-Europeans are not North-Eurasians, but Uralics are. However, the Uralic and Indo-European languages do show remarkable similarities. While some linguists consider that a genetic connection others don't. Most seem to agree though that these similarities mean they "grew up" next door to each other. Could it be that the ANE admixture may be a signal for Indo-Europeans, but that the actual IE carried only a part ANE to Europe? I am merely speculating - I am also merely a layman - but since Belorussinas and Ukranians also carry quite a large part of the WHG admixture I could imagine that parts of West-European WHG admixture was actually brought by Indo-Europeans. Just a thought though.

N haplo is not European, they are mongols that came later and raped some blond women.

 

[LeBrok]

N haplo is not European, they are mongols that came later and raped some blond women.

Please Idun, keep your phobias in check.

 

[ElHorsto]

On fennoscandia they have an interesting comparison:

http://fennoscandia.blogspot.no/2013/09/la-brana-1-and-modern-european-variation.html
http://fennoscandia.blogspot.no/2013/09/la-brana-2-and-modern-european-variation.html

La Brana 2 clusters with Swedes there.

EDIT: With Sami shown separate from Swedes, that is.

It is pretty sure that Swedes are admixed with neolithic farmers to some extent (20%-25% in K12b and similar runs, but also consistently in the large admixture runs by the authors of this paper). The Saami are lacking this admixture completely.
So I wonder if this similarity of La Brana 2 with Swedes could be merely due to the southern admixture, resulting in similar admixture composition, because La Brana showed significant southern admixture in K12b for instance. The PWC hunter-gatherers ajv52/70 found in Gotland had almost no southern admixture and they clustered with Saami and Finns.
To me it is still unclear whether some hunter-gatherers in Iberia already experienced southern admixture (e.g. by farmers or near-eastern hunter-gatherers) or not. The authors rather suggest that the farmers were the ones which are admixed with hunter-gatherers, but I still think this is only half of the truth.

 

[LeBrok]

I could imagine that parts of West-European WHG admixture was actually brought by Indo-Europeans. Just a thought though.

I think that at 7000 years ago WHG was all over the Europe already, it is called west because of Iberian La Brana find, and could originated there during LGM period. 7k ago is too early for IE to spread WHG around.

On page 6 of the Preview PDF, the authors state that Loschbour clusters with 7000 year old hunter-gatherers from Spain, allowing the authors to propose a "West European Hunter-Gatherer (WHG) meta-population. The footnote referenced makes it clear that the authors were referring to the La Brana finds. So, WHG = whoever those two mesolithic people at La Brana were.

 

[epoch]

I think that at 7000 years ago WHG was all over the Europe already, it is called west because of Iberian La Brana find, and could originated there during LGM period. 7k ago is too early for IE to spread WHG around.

But perhaps WHG was even further eastward than all over Europe. Mal'ta showed it partialy. So where ever IE originated, part WHG was then among them.

Otherwise, where is the mesolithic H1 and H3 to make up for the missing amount of mtDNA to equal the 43% WHG admixture in the Norwegians? Or am I missing something?

EDIT: This link: http://biorxiv.org/content/biorxiv/suppl/2013/12/23/001552.DC1/001552-2.pdf

 

[ElHorsto]

http://biorxiv.org/content/biorxiv/suppl/2013/12/23/001552.DC1/001552-1.pdf

The skull of the Loschbourg Hunter-Gatherer looks round and small and the browridges are huge, very strange (Fig. 1A). The La Brana skulls also were not Cro-Magnon-like I think. Don't know what it means. If any anthro-skilled members could comment this, it would be nice.

 

[LeBrok]

But perhaps WHG was even further eastward than all over Europe. Mal'ta showed it partialy. So where ever IE originated, part WHG was then among them.

Otherwise, where is the mesolithic H1 and H3 to make up for the missing amount of mtDNA to equal the 43% WHG admixture in the Norwegians? Or am I missing something?

EDIT: This link: http://biorxiv.org/content/biorxiv/suppl/2013/12/23/001552.DC1/001552-2.pdf

Certainly there was some WHG in IE, it is just hard to say how much at the moment. This WHG might have been stronger in R1a who came to Scandinavia with Corded Ware than with R1b to Western Europe.

Otherwise, where is the mesolithic H1 and H3 to make up for the missing amount of mtDNA to equal the 43% WHG admixture in the Norwegians? Or am I missing something?

the Mt haplogroups don't need to necessary add up to confirm 43% autosomal admixture. It is nice if they do, but they don't contradict autosomal findings.

 

[sparkey]

True, but where did I1 come from if not Mesolithic Scandinavia ? I think it is still a sample bias and that I1 or pre-I1 will definitely show up in other Mesolithic Scandinavian samples.

I think there is growing consensus that modern I1 has higher diversity in Northern Germany than in Scandinavia, and perhaps higher diversity yet in Pomerania than in Northern Germany. My best guess is that it was a small clade somewhere around the South Baltic, and got swept up in a westward expanding population of some sort (Corded Ware or something).

Ken Nordtvedt and others were wrong in their age estimation of I2a1b. It is much older than they thought.

No proof of that yet though. These results actually fit nicely in the Nordtvedt tree, and in fact seem to help fill in the apparent gap in I2a1b's modern distribution, which has always been split oddly between the far northwest of Europe and Eastern Europe.

I have criticised many times (eg here) the method for estimating the age of TMRCA of haplogroups, explaining that most estimates failed to take into account the vastly different historical population sizes of various geographic regions . Warm and fertile countries that adopted agriculture early had much bigger historical populations than northern regions where the hunter-gathering lifestyle survived much longer. Small populations produce less mutations and therefore their haplogroups appear younger than they really are. That is why the age of I subclades is constantly underestimated in relation to Neolithic lineages. This is also true for the mtDNA of hunter-gatherers (see here and here).

I could understand a variable average generation time, and perhaps even climate-influenced mutation rates (although I don't know of any evidence for that...), but if true, how would that apply to I2a-Din, which had a relatively southern expansion some time after the adoption of farming? It would only apply to ancestral I2a1b, and I don't think the exact branching time of I2a1b with I2a1-L1294 has many implications with respect to the spread of I2a-Din, or of our finding I2a1b* in Mesolithic samples.

 

[epoch]

the Mt haplogroups don't need to necessary add up to confirm 43% autosomal admixture. It is nice if they do, but they don't contradict autosomal findings.

Yes, I do realize that. Especially since mtDNA and Y-DNA are far more sensitive to genetic drift: The mother with four sons will loose her mtDNA line, yet contribute largely to autosomal DNA in a polygamous society if her sons take a number of wives, to make a point. But the pattern is so widespread. Almost all of Northwest Europe show little U5 and high H1 + H3. In quite a similar way.

 

[ElHorsto]

But perhaps WHG was even further eastward than all over Europe. Mal'ta showed it partialy. So where ever IE originated, part WHG was then among them.

Otherwise, where is the mesolithic H1 and H3 to make up for the missing amount of mtDNA to equal the 43% WHG admixture in the Norwegians? Or am I missing something?

EDIT: This link: http://biorxiv.org/content/biorxiv/suppl/2013/12/23/001552.DC1/001552-2.pdf

The evidence of paleolithic or mesolithic H is so sparse that I wonder whether it is real:

http://www.ancestraljourneys.org/palaeolithicdna.shtml

http://www.ancestraljourneys.org/mesolithicdna.shtml

But even if it is real for west europe, it must have hibernated somewhere there hidden for a long time and suddenly exploded during Bronze age. The problem is that H multiplied suddenly so much while U remained quite stable in frequency. So even if H is paleolithic, it must be special. I would not discount the possibility that most of it appeared by Bell-Beakers and/or R1b newcomers.

 

[Tabaccus Maximus]

Several things are becoming increasingly clear about H1 and H3:

1. If they did exist in Spain prior to the LGM, they apparently did not expand from there for many thousands of years prior to the Bronze Age. This is a hard sell, especially in light of La Brana which was also U5 and where La Brana-esque people did populate Europe. As resolution of European Mesolithic haplotypes improves, the supposed finds of Paleolithic and Mesolithic H becomes more questionable, given 1) they are generally older studies and 2) proper contextual identification (ie. Paglicci Cave shenanigans).

2. H1 has its highest diversity/frequency in the Syrian highlands and the North Caucacus. It is additionally found at respectable levels in Eastern Europe and among Turkoid peoples in Siberia. Doesn't sound Franco-Cantabrian to me.

3. H1 and H3 are both barely old enough to have experienced the LGM, much less trek to a distant place by some unknown culture, that apparently wasn't Gravettian, to hunker down before the snow flakes started blowing in LGM Europe. If later, then that would defeat the whole purpose of a "refuge hypothesis", which I think is bogus anyway.

4. The diversity, coalescent age and movement of H1 and H3 almost matches exactly the movement of M-269 from Upper Mespotamia/Syria/Anatolia via the North Caucasus into Yamnaya and from there Eastern Europe and Eastern Siberia. It also matches the origin and movement of the Secondary Products Revolution where lactose tolerance is near universal in places where these clades of H are found.

A believable scenario is that H subclades began moving into Europe with the earliest farming where they are found at reduced frequencies. Beaker people brought H1 and H3 from somewhere in the Near East to Spain, where they expanded from there, albeit unevenly, as the Kromsdorf men appear to have taken local women in Central Europe who were the maternal combinations of all previous ages (U2, U5, T1, K, W5, I1, etc)

 

[Maciamo]

2. H1 has its highest diversity/frequency in the Syrian highlands and the North Caucacus. It is additionally found at respectable levels in Eastern Europe and among Turkoid peoples in Siberia. Doesn't sound Franco-Cantabrian to me.

Where did you get that info ? There are very few mtDNA studies on Syria, and all the data I have shows 0% of H1 or H3 in Syria.

3. H1 and H3 are both barely old enough to have experienced the LGM, much less trek to a distant place by some unknown culture, that apparently wasn't Gravettian, to hunker down before the snow flakes started blowing in LGM Europe. If later, then that would defeat the whole purpose of a "refuge hypothesis", which I think is bogus anyway.

I have criticised many times (e.g. here, here and here) the method for estimating the age of haplogroups, explaining that most estimates failed to take into account the vastly different historical population sizes of various geographic regions . Warm and fertile countries that adopted agriculture early had much bigger historical populations than northern regions where the hunter-gathering lifestyle survived much longer. Small populations produce less mutations and therefore their haplogroups appear younger than they really are. That is why the age of I subclades is constantly underestimated in relation to Neolithic lineages. This is also true for the mtDNA of hunter-gatherers.

4. The diversity, coalescent age and movement of H1 and H3 almost matches exactly the movement of M-269 from Upper Mespotamia/Syria/Anatolia via the North Caucasus into Yamnaya and from there Eastern Europe and Eastern Siberia. It also matches the origin and movement of the Secondary Products Revolution where lactose tolerance is near universal in places where these clades of H are found.

No it doesn't since H3 is virtually absent from the Near East and Caucasus, and H1 is pretty rare there too compared to Western and Northern Europe. I have many arguments to disprove that the Indo-Europeans (R1a or R1b) spread H1 and/or H3:

1) H1 and H3 reach one of their highest frequency in Sardinia (31.5%), which is the least Indo-European region in all Europe. Sardinia has one of the lowest percentages of haplogroup R1, close to 0% of East European admixture in the Dodecad, and hardly any ANE in this new study. The only R1b in Sardinia dates from the last 2000 years, notably the Roman and Catalan colonisations.

A believable scenario is that H subclades began moving into Europe with the earliest farming where they are found at reduced frequencies. Beaker people brought H1 and H3 from somewhere in the Near East to Spain, where they expanded from there, albeit unevenly, as the Kromsdorf men appear to have taken local women in Central Europe who were the maternal combinations of all previous ages (U2, U5, T1, K, W5, I1, etc)

It is a terrible mistake to lump all H subclades together as belonging either to Mesolithic European or to later Near Eastern arrivals (Neolithic or Bronze Age). Haplogroup H was present both in Europe and the Near East since the Palaeolithic. One of the oldest identified H sample is actually from Russia. H has also been found in Palaeolithic and Mesolithic Iberia.

I agree that the Beaker expansion could have spread H1 and H3 around Western Europe, but it was surely already present from earlier (Mesolithic & Megalithic) migrations. It just increased their frequency.

 

[Maciamo]

I think there is growing consensus that modern I1 has higher diversity in Northern Germany than in Scandinavia, and perhaps higher diversity yet in Pomerania than in Northern Germany. My best guess is that it was a small clade somewhere around the South Baltic, and got swept up in a westward expanding population of some sort (Corded Ware or something).

That's a possibility. It's just a bit hard to imagine how I1 remained isolated from the numerous LBK-related Neolithic cultures in Germany and Poland for nearly three millennia (c. 5500 to 2800 BCE), then that the R1a expansion of the Corded Ware culture picked up almost exclusively I1 and I2a2 lineages with them to Scandinavia, while the majority of Neolithic lineages in the region must have been Near Eastern (E, G, J, T). Even if I1 and I2a2 did maintain their hunter-gathering lifestyle, living in completely secluded communities side-by-side with the Neolithic farmers, how could I1 lineages have become so numerous in comparison to all other haplogroups, even exceeding the R1a of the Chalcolithic invaders at a ratio of 2:1 ? It doesn't make sense.

If the population of northern Germany and Pomerania had been swept up in the expansion of the Corded Ware culture, we would expect a population in which R1a is dominant, and the rest being a blend of various Mesolithic (I1, I2) and Neolithic lineages (E, G, J, T), but with surely a larger Neolithic faction since farmers could keep much larger populations than hunter-gatherers. I can think of three other possibilities, but all pretty far-fetched:

1) I1 hunter-gatherers of northern Germany adopted agriculture on their own by copying their LBK neighbours, or absorbing a tiny number of LBK lineages. I1 farmers then spread agriculture to Scandinavia, where hunter-gatherers belonged to I2a1 (Motala). Since the autosomal DNA of Gök4, a Neolithic farmer from Sweden, was clearly Near Eastern, it renders the possibility of I1 introducing agriculture extremely unlikely.

2) I1 was part of the original Corded Ware people alongside R1a, or was picked up very early on around Belarus and eastern Poland. Through a dramatic founder effect, the Corded Ware people who settled in Scandinavia were mostly I1, with a R1a and I2a2 minority. However I1 has never shown up in ancient sites in Eastern Europe, nor in the Corded Ware samples from Eulau, which contained only R1a. The likelihood is therefore also low.

3) Corded Ware was truly an R1a-dominated expansion. LBK lineages dominated most of Germany and Poland, except in the northern coastal areas, which were almost exclusively inhabited by I1 hunter-gatherers. When R1a arrived along the Baltic coast, I1 people quickly adopted their ways of life and technologies, and the Corded Ware was spread from northern Germany and Denmark to Sweden and Norway by I1 and R1a people. There is no evidence supporting such a far-fetched scenario either.

That is why I think that the most likely scenario is that I1 was the lineage of Mesolithic Scandinavians all along, and that the I2* and I2a1b samples from Motala are simply unrepresentative. Two other Motala samples (#2 and #9) were described as I*, but could very well have been pre-I1.

No proof of that yet though. These results actually fit nicely in the Nordtvedt tree, and in fact seem to help fill in the apparent gap in I2a1b's modern distribution, which has always been split oddly between the far northwest of Europe and Eastern Europe.

I might have had some older version of Nordtvedt tree in mind. It's true that the current tree fits.

I could understand a variable average generation time, and perhaps even climate-influenced mutation rates (although I don't know of any evidence for that...), but if true, how would that apply to I2a-Din, which had a relatively southern expansion some time after the adoption of farming? It would only apply to ancestral I2a1b, and I don't think the exact branching time of I2a1b with I2a1-L1294 has many implications with respect to the spread of I2a-Din, or of our finding I2a1b* in Mesolithic samples.

If I2a-Din had a relatively southern expansion some time after the adoption of farming, then it makes more sense that I2a-Din spread relatively early, first with the Cucuteni-Tripillian culture (4800-3000 BCE), then with the Proto-Thracians of the Multi-cordoned ware culture (2200-1800 BCE), and eventually with the Daco-Thracians and Illyrians, rather than with the Slavs several millennia later. Wouldn't it ?

 

[Idun]

Please Idun, keep your phobias in check.

Just having fun as you try to work around this stuff.