Massive migration from the steppe is a source for Indo-European languages in Europe

[Goga]

I have proposed many years ago that the J2a of Indian Brahmins was picked up in southern central Asia when R1a Proto-Indo-Iranians mixed with the local population in the BMAC complex, before invading India and Iran. That would explain why the Balto-Slavic R1a branch, or northern European R1b for that matter, do not have any meaningful percentage of J2a.

I don't agree with you. (proto-)Iranic is closer to Greek than to Balto-Slavic, the Satem link between Iranic and Balto-Slavic is not important here. Origin of the Iranic languages must be very different than the origin of Balto-Slavic. Not only Indo-Iranic (Iranians and Indians) people have J2a, but also Anatolian Indo-Europeans, like Armenians, and Greeks have J2a in them too. Basically more than 20-30% of all modern day Indo-Europeans speakers (mostly who live outside Europe) carry this haplogroup. Also it's one of the most widespread haplogroups among the Indo-European speakers, from NorthWest Europe to SouthEast Asia (India) and has high distribution in Northern Caucasus, Maykop Horizon..

 

[Tone]

Give me a break, please:

• Why R1b couldn't have been spread around Western Europe by the Bell Beaker people
• Bell Beakers were a multicultural phenomenon & trade network, not an ethnic culture

Sorry. I didn't mean to offend you by asking questions. I'll go now.

 

[bicicleur 2]

Then it means that the original Anatolian branch wasn't Z2103, but P297*, M269* or L23*. Perhaps it is the Cimmerians, the Sarmatians or even the European Scythians who brought Z2103 to the Balkans and Anatolia much later.

that is the most likely explanation

3300-2600 BC, it is before Sintashta
Indo-Iranians went east and south with chariots
somebody must have brought chariots to the Balkans

 

[Sile]

I know that some Finns will get upset about this,but check this interesting theory about the name of Sami which could also be related to the name of Finland:
http://en.wikipedia.org/wiki/Sami_people#Etymologies
...
"The first known historical mention of the Sami, naming them Fenni, was by Tacitus, about 98 A.D.^[13] Variants of Finn or Fenni were in wide use in ancient times, judging from the names Fenni and Phinnoi in classical Roman and Greek works. Finn (or variants, such as skridfinn, "striding Finn") was the name originally used by Norse speakers (and their proto-Norse speaking ancestors) to refer to the Sami, as attested in the Icelandic Eddas and Norse sagas (11th to 14th centuries). The etymology is somewhat uncertain, but the consensus seems to be that it is related to Old Norse finna, from proto-Germanic *finthanan ("to find"),^[14] the logic being that the Sami, as hunter-gatherers "found" their food, rather than grew it. It has been suggested, however, that it may originally have been a more general term for "northern hunter gatherers", rather than referring exclusively to the Sami, which may explain why two Swedish runestones from the 11th century apparently refer to what is now southwestern Finland as Finland. Note that in Finnish, Finns (inhabitants of Finland) do not refer to themselves as Finns. As Old Norse gradually developed into the separate Scandinavian languages, Swedes apparently took to using Finn exclusively to refer to inhabitants of Finland, while Sami came to be called Lapps. In Norway, however, Sami were still called Finns at least until the modern era (reflected in toponyms like Finnmark, Finnsnes, Finnfjord and Finnøy) and some Northern Norwegians will still occasionally use Finn to refer to Sami people, although the Sami themselves now consider this to be a pejorative term."
.....
According to the history,Vikings were not only raiding,but they were raising animals (goats,pigs,etc) ,fishing and even practice agriculture.
Think that Scandinavia is good example of how Indo-European speaking population interacted with native population,here we have Northern Germans (Vikings) interacting with native Fino-Ugric people,which included also Sami people.

Here some archeological proof,that Vikings were practicing agriculture 1000 years ago,even in Greenland:
http://sciencenordic.com/vikings-grew-barley-greenland

maybe you need to go a bit further back and read this first

http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2986642/

 

[mihaitzateo]

maybe you need to go a bit further back and read this first

http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2986642/

Have read it,but I do not see anything told about the Finns autosomal admixture,which should have significant more Hunter-Gatherer admixture,versus Swedish admixture,which should have lots of Yamnaya with more Neolithic.

Anyway,a very strange thing that Norwegians are clustering most close to Yamnaya and I do not know what Norwegians are those tested,I think if you test Norwegians from near sea side,from South Norway,those will cluster even more closed to Yamnaya having fewer HG admixture.
Another question is when Yamnaya people started to move from there towards Western Europe.
I understand that it dates from around 3600-2300 BC.
Maybe in those times Celts and Germans and Italics were speaking same language and as they moved from there,their languages split.
Is a possibility that Italics moved from there by boat,and got into Italy,while Celtic and Germanics traveling on the ground.

 

[Angela]

Yes, great results. Never expected such a confirmation of my theories. No R1a in Yamnaya, so there's still no evidence that R1a-Z93 in Iranic folks is from Yamnaya or the Pontic-Caspian Steppes in general. I knew it, but didn't expect that they would find Anatolian (Armenian, West Iranic) R1b in Yamnaya. R1b in Yamnaya is Anatolia, which again is a great indication that Vyacheslav Vsevolodovich Ivanov is right about his Armenian hypothesis. http://en.wikipedia.org/wiki/Armenian_hypothesis . The latest results are victory for Vyacheslav Vsevolodovich Ivanov & Tamaz V. Gamkrelidze! It's true that Indo-Europeans in Europe came from Yamnaya. But folks from NorthWest Asia (from Maykop) Indo-Europized the Yamnaya folks in the Steppes. I was telling this all the time. Indo-Europeanization occured in stages. Best news for me is that R1a-Z93 has nothing to do with the Yamnaya. And this fact is making my thoughts even stronger!

Goga, I think you're jumping the gun a bit if you don't mind me saying so in terms of the actual genetic proof for these things.

The oldest basal R1b is a hunter gatherer from Samara, grouped within EHG, along with the R1a Karelian. There's nothing "Near Eastern" about him.

"The hunter-gatherer from Samara belonged to haplogroup R1b1 (L278:18914441C→T), with
upstream haplogroup R1b (M343:2887824C→A) also supported. However, he was ancestral for both
the downstream haplogroup R1b1a1 (M478:23444054T→C) and R1b1a2 (M269:22739367T→C) and
could be designated as R1b1*(xR1b1a1, R1b1a2). Thus, this individual was basal to most west
Eurasian R1b individuals which belong to the R-M269 lineage as well as to the related R-M73/M478
lineage that has a predominantly non-European distribution."

His culture included ceramics but there was no pastoralism at that time in that area. The Yamnaya samples were from one specific site-Samara, and one of them is one is, and five are It is only these samples which are half "Armenian-like".

At present, the only way I can see for the men of these particular subclades to have come from the Caucasus or south of it bearing Near Eastern ancestry is if R1b was a hunter gatherer lineage which existed both north of the Caucasus mountains and in them (and perhaps a little south of them as well, and came into contact with more "southern" populations.) However, this Samara R1b man was half WHG. How much WHG is present south of the Caucasus? I don't think very much, but I don't have the data right at my fingertips so perhaps someone can provide it. Also, what about within the Caucasus? Of course, we now know modern populations are not good proxies for ancient autosomal components, so there is that to consider, and there's been a lot of traffic in that area. Also, I suppose it's possible that the R1b Samara man got his WHG from mixing with Karelia like people?

Another thing in support of this theory it seems to me is, strangely enough, that R1b1 Neolithic sample in Spain. If R1b when it was still hunter gatherer was more ANE than anything else, was all over the Caucasus area, then it's understandable how some of them were far south of the Caucasus enough to be swept along by the Neolithic and wind up in Spain. Then, the downstream clades could have developed south of the Caucasus and gone back up north as well.


The other possibility is that the downstream clades developed right on the Steppe (never having moved at all),and the Near Eastern like ancestry came either from men (and the women accompanying them) in areas of the Yamnaya horizon not yet sampled (there is that J2a1 Bronze Age Indo-European in central Europe to consider), or it comes only from women.

I discussed this a bit with Greying Wanderer in connection to 6,000 BC interactions. I still think it's a bit of a stretch to see hunter-gatherers males going all the way to the South Caucasus to raid women, but we do have Maykop right on the brink of the Steppes, and a strong argument can be made, I think, that some of the metal working and other parts of the "Yamnya" package came from them. Given the kind of host/patronage relationships characteristic of the Indo-Europeans (and their strongly patriarchal nature), perhaps the women were given to the Steppe men as part of some type of exchange. The bartering of women to cement trade and other economic and political partnerships was part of royal and aristocratic mating virtually down to the present time.

Anyway, until we have more samples from all parts of the Yamnaya horizon and south into the Caucasus, I don't think we can be much more definite than this. Well, I can't be more definite than this at the present time.

(I just realized that this may have been already addressed. Maybe I should start from the end of the thread and work backwards. )

 

[Sile]

Note the chart on page 23...zero WHG for tuscans ..............what does that do for tuscan admixture

 

[Maciamo]

Sorry. I didn't mean to offend you by asking questions. I'll go now.

Don't worry, I am not offended. It's true that you are a relatively new member and we have been discussing the Bell Beaker issue for several years on this forum.

 

[Maciamo]

I must admit I have not that much knowledge about Bell Beakers thats why the question mark.

But than the chart shows late Bell Beaker was ~50% Yamna like.

Late Beakers in Germany. I am very confident that it wasn't the case at all in Iberia, where Yamna influence was minimal (although probably not entirely absent in some regions after 2000 BCE).

 

[Angela]

As to what happened to some of the WHG figures in Europe, perhaps this chart would be helpful:

 

[Sile]

Note the chart on page 23...zero WHG for tuscans ..............what does that do for tuscan admixture

A quick scenario by me, would indicate that etruscan main haplogroup origin is not R1b-U152 but J2

R1b-U152 would be clearly a celtic - gallic -liguri marker

 

[Greying Wanderer]

Give me a break, please:

• Why R1b couldn't have been spread around Western Europe by the Bell Beaker people
• Bell Beakers were a multicultural phenomenon & trade network, not an ethnic culture

Copper smiths and miners spreading along the neolithic trade network with some of them settling as artisans among many different cultures including the Atlantic Megalith culture and then setting up mining colonies from that base along the under populated Atlantic coast regions is why they could have spread R1b to specifically the Atlantic coast part of western Europe *separate from* the main source of R1b coming overland.

They would have needed some extra ingredient to explain how a small population could dramatically expand in those under populated regions when LBK couldn't. I wonder what that could be.

LBK map (note gap to west and north of LBK range)

http://what-when-how.com/wp-content/uploads/2011/03/tmp3517_thumb1.jpg

LP map (note peaks along those same gaps)

http://1.bp.blogspot.com/-k4oPZEUau...encies+based+on+all+genotype+frequencies..png

 

[Aberdeen]

Norwegians have a higher combined percentage of R1a + R1b than Belarussians and Ukrainians. Plenty of Central Asians invaded eastern Europe over the last 5000 years, almost completely eliminating R1b in the region. I explained 5 years ago that this was why R1b was so low today in its original homeland.

The huge Neolithic population of the Cucuteni-Trypillian culture didn't just vanish in thin air. They were gradually absorbed by PIE people (probably already since the Globular Amphora culture). Don't forget that Cucuteni-Trypillian towns were the largest in the world at the time. That explains the very significant percentage of both male and female Near Eastern lineages in western Ukraine and southern Belarus today.

Additionally, Ukrainians also have partial Greek ancestry in the south (lots of J2a).

It is especially northern Belarus and eastern Ukraine that are very high in R1a, and that is just a sign of higher recent Slavic ancestry, not a sign of more surviving Yamna ancestry. The Slavic branch descends from the Corded Ware and Abashevo cultures, not from Yamna.

Actually, according to your own tables, Norwegians don't have that much more R1a + R1b than Belarusians or Ukranians. What they do have is a lot more I1 and I2 haplotype material than Belarusians or Ukranians. So, although I know that Y haplotype frequency doesn't necessarily tell us about autosomal content, I would expect Norwegians with all that I1 and I2b to be fairly high in WHG, higher in fact than Belarusians and Ukrainians. Perhaps there's a reason for Nowegians apparently having minimal WHG in spite of having 31.5 I1 and 4.5 I2b, but I suspect the reason is that the calculation method doesn't distinguish properly between WHG and Yamnaya.

 

[mihaitzateo]

Note the chart on page 23...zero WHG for tuscans ..............what does that do for tuscan admixture

I am supposing that Tuscans are most closed from Ancient Romans,from populations sampled there.

Is interesting the legend of Europe which is described in old Greek legends,Zeus is seducing a maiden from an island ,the maiden being called Europe and being of Phoenician ancestry.
http://en.wikipedia.org/wiki/Europa_(mythology)
Europa is an Middle East woman,in the legend.
And Zeus come as a white bull.
So the supposition is that Zeus is representing the R1/R1B people who are coming and conquering the old farmers (Early Neolithic) people from Europe.
Is just a supposition.
Look at the Greeks,they are having lots of Early Farmers admixture.
And we can suppose old Greeks were even higher on Early Neolithic admixture (farmers) and lower on Hunter Gatherer and Yamnaya,which were brought by mixing with Slavs,with some Goths ,that settled in Greece and so on.
It should also be noticed that in Greece we have highest percentage of J2 so is quite clear I think that Greeks between Europeans have most from Early Neolithic people,the farmers people.
And R1A and R1B are representing the barbarians.
(which are represented by Zeus and other Olimp deities,probably an Old Greeks irony at the fact that lots of Indo-Europeans had in their religion the belief they are descending from gods ).

Wonder how Ancient Romans admixture was,I am inclined to believe that Ancient Romans were mostly J2 also and since lots died in wars they replaced with assimilated Gaulish-Celtic people the lost people,this is why R1B is at such high rates in Italy.

 

[Maciamo]

True but ancient samples have taught us that allot of our understanding of DNA can not be explained with it's modern distribution. The same way with Z2103. If we would have used it's distrbution in modern Europe or South_Central Asia as argument wether it is Indo European or not. We would have drifted completely into the wrong direction.

I do not think J2a was simply picked up by Indo Iranians in Central Asia since the Bronze Age J2a in Hungary doesn't seem very Indo_Iranian to me. Also a Neolithic origin can be excluded. To be honest only an Indo European explanation for it's origin remains.

I wouldn't read too much into the Late Bronze Age J2a1 sample from Hungary (c. 1200 BCE). The Indo-Europeans had been in the region for nearly 2000 years by then, and it could very well have been a foreigner or an assimilated person from a non-IE culture. After all three Unetice samples tested in this new study turned out to be assimilated local I2 lineages, not steppe lineages.

We are talking about one of the most unstable period in European and Near Eastern history here. Most East Mediterranean civilizations collapsed around 1200 BCE due to the mysterious Sea Peoples. Even the Haak el al. paper discussed here mentions that 1200 BCE is the turning point for the domination of Yamna-like admixture in central European graves. As I mentioned above, this also happen to correspond to the sudden adoption of cremation instead of kurgan/tumulus burial in central Europe for only one hundred years (Urnfield culture), before the traditional tumulus burial resumed with the Hallstatt culture.

No one knows precisely what cataclysmic events took place around 1200 BCE, but I would bet that this corresponds to an expansion of J2a people in the Eastern Mediterranean, and that the Sea Peoples were probably predominantly J2a people. After all, all the great ancient seafaring civilizations all presumably had high percentages of J2a, including the Phoenicians who just happen to emerge around 1200 BCE.

One hypothesis of mine is that the Sea Peoples were descended from the Minoan civilization, which had just collapsed c. 1450 BCE. The fall of the Minoan state might have led to Minoan people turning to piracy and raiding the coasts of the East Mediterranean, from Sardinia to the Levant. The Trojans, whose city had been destroyed by the Mycenaeans c. 1200 BCE, may well have been a mixture of R1b-L23 and J2a people, and Trojans who escaped may also have been among the Sea Peoples who sought revenge on Mycenaean Greece and caused its downfall less than 100 years later.

1200 BCE also coincides with the arrival of the Dorians in Greece, the Phrygians in western Anatolia and the Armenians in eastern Anatolia, who I think were all predominantly R1b-L23 tribes from the Balkans, with assimilated Neolithic lineages (E-V13, G2a, I2...). Their departure from the Balkans left place for a northward expansion of Greek lineages, among which J2a.

In such a context, it would be unreasonable to label a Hungarian J2a1 as necessarily Indo-European.

 

[Greying Wanderer]

@Angela

I still think it's a bit of a stretch to see hunter-gatherers males going all the way to the South Caucasus to raid women

Yes, a follow-on of that theory is (edit: it implies) the "Armenian-like" population may have moved south to get away from them i.e. they were originally in the north and close enough to be raided (or traded with) earlier.

 

[mihaitzateo]

Lol Maciamo are you serious when you are saying that R1b were displaced from Eastern Europe by raiding Central Asians?

I rather think that somehow not too many R1B people do not got into the area of South Eastern Europe. And the area was already very likely quite populated with Neolithic farmers.
For example,we are seeing lots of E-V13 and J2 and I2 in Balkans and even Romania,well the area had dense forests,lots of mountains,it was hard to get there,from where Yamna home was.
A very interesting thing would be to get at least a clue about how the forests were spread in those times,in this part of Europe.

 

[Yetos]

I wouldn't read too much into the Late Bronze Age J2a1 sample from Hungary (c. 1200 BCE). The Indo-Europeans had been in the region for nearly 2000 years by then, and it could very well have been a foreigner or an assimilated person from a non-IE culture. After all three Unetice samples tested in this new study turned out to be assimilated local I2 lineages, not steppe lineages.

We are talking about one of the most unstable period in European and Near Eastern history here. Most East Mediterranean civilizations collapsed around 1200 BCE due to the mysterious Sea Peoples. Even the Haak el al. paper discussed here mentions that 1200 BCE is the turning point for the domination of Yamna-like admixture in central European graves. As I mentioned above, this also happen to correspond to the sudden adoption of cremation instead of kurgan/tumulus burial in central Europe for only one hundred years (Urnfield culture), before the traditional tumulus burial resumed with the Hallstatt culture.

No one knows precisely what cataclysmic events took place around 1200 BCE, but I would bet that this corresponds to an expansion of J2a people in the Eastern Mediterranean, and that the Sea Peoples were probably predominantly J2a people. After all, all the great ancient seafaring civilizations all presumably had high percentages of J2a, including the Phoenicians who just happen to emerge around 1200 BCE.

One hypothesis of mine is that the Sea Peoples were descended from the Minoan civilization, which had just collapsed c. 1450 BCE. The fall of the Minoan state might have led to Minoan people turning to piracy and raiding the coasts of the East Mediterranean, from Sardinia to the Levant. The Trojans, whose city had been destroyed by the Mycenaeans c. 1200 BCE, may well have been a mixture of R1b-L23 and J2a people, and Trojans who escaped may also have been among the Sea Peoples who sought revenge on Mycenaean Greece and caused its downfall less than 100 years later.

1200 BCE also coincides with the arrival of the Dorians in Greece, the Phrygians in western Anatolia and the Armenians in eastern Anatolia, who I think were all predominantly R1b-L23 tribes from the Balkans, with assimilated Neolithic lineages (E-V13, G2a, I2...). Their departure from the Balkans left place for a northward expansion of Greek lineages, among which J2a.

In such a context, it would be unreasonable to label a Hungarian J2a1 as necessarily Indo-European.

hmm

in such case , Varna necropolis, Rudna Glava, and generally Vinca and para-Vinca cultures had no J2a,
I mean if J2a is upsent from Balkans before 1200 BC, they should be upsent also from copper mettalurgy, not bronze, but copper,
in such case who brought copper mettalurgy to balkans?
we know Vinca and para-Vincas are before bronze, meaning before R1*****, and Mycenae where build before 4000 BC 1500 years before arsenic bronze from steppe, theoritically start to enter south Balkans
beside in many settlements in Greece as the newly Platamon metaneolithic we see no arsenic bronze, in many areas till even after 1500 BC.
but Vincas knew copper and gold very well.


so then if J2xyz did not exist in Vincas, neither R1xyz, could copper spread there by? G2a? Ixyz, ?

 

[Goga]

I wouldn't read too much into the Late Bronze Age J2a1 sample from Hungary (c. 1200 BCE). The Indo-Europeans had been in the region for nearly 2000 years by then, and it could very well have been a foreigner or an assimilated person from a non-IE culture. After all three Unetice samples tested in this new study turned out to be assimilated local I2 lineages, not steppe lineages.

We are talking about one of the most unstable period in European and Near Eastern history here. Most East Mediterranean civilizations collapsed around 1200 BCE due to the mysterious Sea Peoples. Even the Haak el al. paper discussed here mentions that 1200 BCE is the turning point for the domination of Yamna-like admixture in central European graves. As I mentioned above, this also happen to correspond to the sudden adoption of cremation instead of kurgan/tumulus burial in central Europe for only one hundred years (Urnfield culture), before the traditional tumulus burial resumed with the Hallstatt culture.

No one knows precisely what cataclysmic events took place around 1200 BCE, but I would bet that this corresponds to an expansion of J2a people in the Eastern Mediterranean, and that the Sea Peoples were probably predominantly J2a people. After all, all the great ancient seafaring civilizations all presumably had high percentages of J2a, including the Phoenicians who just happen to emerge around 1200 BCE.

One hypothesis of mine is that the Sea Peoples were descended from the Minoan civilization, which had just collapsed c. 1450 BCE. The fall of the Minoan state might have led to Minoan people turning to piracy and raiding the coasts of the East Mediterranean, from Sardinia to the Levant. The Trojans, whose city had been destroyed by the Mycenaeans c. 1200 BCE, may well have been a mixture of R1b-L23 and J2a people, and Trojans who escaped may also have been among the Sea Peoples who sought revenge on Mycenaean Greece and caused its downfall less than 100 years later.

1200 BCE also coincides with the arrival of the Dorians in Greece, the Phrygians in western Anatolia and the Armenians in eastern Anatolia, who I think were all predominantly R1b-L23 tribes from the Balkans, with assimilated Neolithic lineages (E-V13, G2a, I2...). Their departure from the Balkans left place for a northward expansion of Greek lineages, among which J2a.

In such a context, it would be unreasonable to label a Hungarian J2a1 as necessarily Indo-European.

Close, but no cigar. Although it's a very fascinating theory. Nice try, respect! J2a folks were actually MOUNTAIN people and not SEA people. J2a peaks in the Caucasus, Zagros Mountains and SouthCentral Asia. There's lots of ANE and Caucaso-Gedrosian component in the Caucasus region. ANE is highest there in the whole Europe and West Asia. I think that J2a is also correlated with ANE and Caucaso-Gedrosia component. Ancient J2a should be full of ANE and Caucaso-Gedrosian component. What you're talking about is a massive replacement of folks in South Europe around 1200 BCE. I don't think it was the case, because if such a huge migration of so called Sea People with J2a occured in South Europe, there would be much more ANE and Caucaso-Gedrosian component in Southern Europe, even more than in Northern Europe. There's a lot of J2a in modern MAYKOP and it's not from recent times! When R1b migrated from Maykop into Yamnaya, there was already J2a in Maykop!

 

[Angela]

As to what happened to some of the WHG figures in Europe, perhaps this chart would be helpful:
View attachment 7071

Sorry it's not bigger. However, I think it's worth giving it a look.

I think that these figures have to be unpacked a bit.

What has to be kept in mind is that the EHG figure includes WHG as well as ANE. That is partly why the WHG in modern Europeans in this paper is lower than the WHG in the prior Lazardis et al paper. For example, in this paper the Southwestern French, two-thirds (?) of the Spaniards, the Tuscans, and the Sicilians have 0% WHG, whereas in the prior paper, the Southwestern French were 20% WHG, the Tuscans were 14%, and the Spaniards were 7% WHG.

So, effectively, the WHG in this chart might be the amount of native WHG left in these areas before some new WHG was brought in by the Yamnaya people.

This would explain why the authors say there was a near 100% wipe out of hunter gatherers in some areas by the Neolithic farmers. It would also indicate that there was probably no "resurgence" in southern Europe of the WHG component, which makes it more likely, in my opinion, that, as I had speculated before, the "resurgence" in northern and central Europe is probably the result of hunter gatherers from the peripheral refuges moving south during a climate worsening event, or at least of the additional admixture resulting from contacts along that northern border where the farmers had not penetrated.

Of course, the proportion of WHG (and not some UHG from the Near East) in the early neolithic farmers should be added to this new number as well, yes?

The EN figure probably includes, therefore, not just the EEF like people but also the "Near Eastern" portion of the Yamnaya people.

Ed. Any resurgence in Spain and Northern Italy, given that they have 2-3% WHG of perhaps the "original" variety, might come from WHG who took refuge in the Pyrenees and the Alps.

Also, it should be noted that later gene flows could have further decreased WHG levels. For example, in order to get this close a fit, the authors had to add Nganasans and Bedouins.