# Population Genetics > Paleogenetics > Paleolithic & Mesolithic >  Revisiting the issue of "Ice Age refugia" in the context of WHG, SHG, EHG ancestries

## Tomenable

We have learned by now, that there were at least three, autosomally quite distinct, types of foragers in Europe:

WHG - western (from Iberia to North-Western France) hunter-gatherers
SHG - central (from Scandinavia at least to Hungary*) hunter-gatherers
EHG - eastern (from Karelia to Pontic-Caspian area) hunter-gatherers

*This is what Pinhasi claimed ("Pinhasi’s team has found that the genomes sequenced from hunter-gatherers from Hungary and Switzerland between 14,000 to 7,500 years ago are very close to specimens from Denmark or Sweden from the same period" - LINK).

There were similarities between them, but also differences. EHG, for instance, was ANE-rich.

Both SHG and EHG were also lighter-pigmented than WHG. WHG were darker, if I remember correctly.

WHG probably had more of Aurignacian ancestry (see Y-DNA of Kostenki14, and then La-Brana).

I think these differences between WHG, SHG and EHG components (combined with lack of significant differences WITHIN each of those components - for example Karelian hunter was autosomally almost identical with Samara hunter) could be the result of their distinct origins from three separate Ice Age refugia. That old concept of Ice Ace Refugia was wrong when it comes to association of certain haplogroups with certain refugia (the map below) - especially when it comes to the spurious association of R1b with the Iberian refugium, on the grounds of modern frequencies.

But it doesn't mean that the entire concept was necessarily erroneous:

https://en.wikipedia.org/wiki/Last_G...aximum_refugia



Association of three HG autosomal components - WHG, SHG and EHG - with those three refugia as shown above, seems probable to me, and could explain observed differences between them. However, when it comes to Y-DNA, WHG were not R1b, but rather C1 (C-M130) and some subclades of I2. The "central", Balkan or Italian, refugium, could be home to other clades of I2 (e.g. those found in Sweden: I2a1, I2c) and to I1.

To sum up, this is my hypothesis:



What do you think about this ???

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If such an explanation of differences between WHG, SHG, EHG was already proposed before, then I apologize for not checking carefully enough.

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## bicicleur

WHG - western (from Iberia to North-Western France) hunter-gatherers
SHG - central (from Scandinavia at least to Hungary*) hunter-gatherers
EHG - eastern (from Karelia to Pontic-Caspian area) hunter-gatherers

where does this concept come from and about what period was this ?

the archeological record hints towards multiple expansions from southern France (Magdalenian, Tardenoisian, Azilian ..)
there was the Italian epigravettian which probably originated in the northern Adriatic Sea, which was dry land during LGM ; this expanded into Italy and the Carpathian Basin, but not any further
there where some ice age HG around the Aegean Sea and along the coasts of Albania and Montenegro, but no expansions from there
there was the eastern Epigravettian probably originated in the northern part of the Black Sea, which was dry land during LGM and expanded into Transcaucasia and till east of the Carpaths (Moldova, eastern Roumenia)
around 9 ka Tardenoisian reached the area east of the Carpaths and some areas inside the Carpathian Basin which were epigravattian territories before that

this whole story does not match the WHG - SHG - EHG division at all

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## Tomenable

> this whole story does not match the WHG - SHG - EHG division at all


So how would you explain this division? BTW - I've also started a similar thread at Anthrogenica:

http://www.anthrogenica.com/showthre...EHG-ancestries

So far I got one response which suggests that SHG was probably just a mixture of WHG and EHG:
(so we are left with just 2 refugia instead of three, if this user is right):




> SHG's likely didn't exist in the Balkans, but are a result of WHG-EHG mixture in Southern Scandinavia. KO1 was just a WHG and early Hungarian neolithic farmers did not have EHG either. Balkan HG's probably were fully WHG.


However, according to Pinhasi, SHG lived also in Hungary and in Switzerland, not just in Southern Scandinavia.

By contrast, in Luxembourg there was already WHG, not SHG. In Iberia there was also WHG.

In Russia there was EHG - Karelian genome and Sok River genome (Samara Oblast) were both of EHG type.

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## Tomenable

Another response:




> Yep, SHG=WHG+EHG seems most likely scenario. There will be same paleolithic DNA from Spain, if this link is true: http://www.newscientist.com/article/...e#.VZ6kNLUtyql


Yet according to prof. Pinhasi (see the link that I posted in the OP), hunters from Hungary and Switzerland were very similar to those from Sweden and Denmark, if I interpreted that correctly. So how was it in the end, were those hunters more similar to those from Iberia and Luxembourg (WHG), or to those from Sweden (SHG)? Do we even have any samples of actual hunters from Hungary and Switzerland?

I think we need samples of hunters from the Balkans and from Italy, to definitely answer this question.

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## ElHorsto

> We have learned by now, that there were at least three, autosomally quite distinct, types of foragers in Europe:
> 
> WHG - western (from Iberia to North-Western France) hunter-gatherers
> SHG - central (from Scandinavia at least to Hungary*) hunter-gatherers
> EHG - eastern (from Karelia to Pontic-Caspian area) hunter-gatherers
> 
> *This is what Pinhasi claimed ("Pinhasi’s team has found that the genomes sequenced from hunter-gatherers from Hungary and Switzerland between 14,000 to 7,500 years ago are very close to specimens from Denmark or Sweden from the same period" - LINK).


Afaik in Gamba et al there was one neolithic sample from Hungary which turned out to be purely WHG on both, ANE-WHG breakdown and PCA plot. He was different from scandinavian hunter-gatherers who had more ANE. So I think SHG is a mixture between WHG and EHG. I don't know why Pinhasi has different results.

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## bicicleur

> So how would you explain this division? BTW - I've also started a similar thread at Anthrogenica:
> 
> http://www.anthrogenica.com/showthre...EHG-ancestries
> 
> So far I got one response which suggests that SHG was probably just a mixture of WHG and EHG:
> (so we are left with just 2 refugia instead of three, if this user is right):
> 
> 
> 
> ...


If the map above is correct , SHG = EHG + WHG then it could be explained as follows :

WHG = I2a , southern France during LGM
EHG = I2b+c, eastern epigravettian , I2b+I2c is the only subclade of I who also has an important presence in the Transcaucasus , so that makes sens

then SHG would be a mixture of I2a and I2b+I2c
hence in Motola 8 ka we find both I2a and I2c2

eastern epigravettian in Transcaucasus lasted till 12 ka , maybe this signals arrival of R1a which was detected in Karelian HG later

just a posibility ..

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## Fire Haired14

2/3 WHGs are I2a1. One was I2a1a(xM26) and one pre-I2a1b. The Scandinavian HGs belonged 100% to I2, and mostly to I2a1. They even shared pre-I2a1b and I2a1a with WHG. Plus, I2 is more popular in Neolithic farmers who lacked ANE than C. So, I2 was probably more popular in WHG than C. 

https://docs.google.com/spreadsheets...oIA/edit#gid=0

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## bicicleur

> 2/3 WHGs are I2a1. One was I2a1a(xM26) and one pre-I2a1b. The Scandinavian HGs belonged 100% to I2, and mostly to I2a1. They even shared pre-I2a1b and I2a1a with WHG. Plus, I2 is more popular in Neolithic farmers who lacked ANE than C. So, I2 was probably more popular in WHG than C. 
> 
> https://docs.google.com/spreadsheets...oIA/edit#gid=0


IMO C1a2-La Brana where Aurignacians, while I were Gravettians
By LGM C1a2 were almost extinct in Europe, only few pockets left. They are no part of the big mesolithic picture in Europe.
C1a2-Neolithic Hungary were descendants of Levantine Aurignacian coming to Europe with G2a, so not part of mesolithic Europe either.
According to YFull TRMCA for C1a2-V20 is 43,2 ka.
First Aurignacians in Wellendorf are dated 43,5 ka , they split from European Mediterranean proto-Aurignacians.
First Levantine Aurignacian is less than 40 ka, IMO they split from European Mediterranean proto-Aurignacians after the Archiflegreo eruption (39,3 ka).
Same for Baradostian, which is categorized as the Zagros Mts Aurignacian.

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## LeBrok

The picture is more muddy. WHG had spread into whole Europe by the end of Ice Age. EHG is a mix between WHG and ANE. We would need to find a place where ANE hid during LGM. Probably few of them with R1a, R1b, C, N and what's not. The mixture of EHG could have happened after Ice Age, so refugium of EHG didn't exist, probably.
SHG developed from mostly WHG, some ANE and independent mutations caused by separation there in Scandinavia for few thousands of years. Again refugium of SHG didn't exist.
We also need to take under consideration that some WHG I2 was hiding in Anatolia.

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## Greying Wanderer

My guess (using the map from rozenfeld above)

SW quadrant = WHG
SE quadrant = ASE
north = archaic or archaic mixed northern interior population

leading to

NW quadrant = WHG + northern archaics = SHG 
NE quadrant = ASE + northern archaics = EHG

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