More DNA from stone age European(Swedish) farmers and hunter gatherers

Fire Haired14

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Y-DNA haplogroup
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mtDNA haplogroup
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Today(April 24, 2014) possibly the greatest ancient DNA paper since Lazaridis 2014 'appeared in Science" Skoglund et al. 2014. I was informed by More ancient Scandinavians (Skoglund, Malmström et al. 2014). I just want to make people aware, i will add my own opinion after reading its Supplementary Materials.

There is already quite a collection of stone age Swedish hunter gatherer and farmer DNA. The individuals from this study are highlighted. If individuals in older studies were sampled from the same site as samples in Skoglund et al. 2014 but were given a differnt date, I replaced the old date with the one given in this paper. Ancestral Journeys.org is my source for the stone age Swedish individuals not in Skoglund et al. 2014


Mesolithic Swedish hunter gatherers
StoraFörvar11 aka SfF11(Male), 7,500-7,250 cal. B.P, Stora Karlso Sweden : mtDNA=U5a1

6,873 ± 119 BC, Stora Karlso Sweden : mtDNA=U4b1

Motala1(Female), 6,000BC Motala Sweden: mtDNA=U5a1

Motala2(Male), 6,000BC Motala Sweden: Y DNA=I* (I P38+, I PF3742+, I L41+, I1 S108-, I1 L845-, I1 M253-, I2a1b CT1293-, I2a2 L37-), mtDNA=U2e1

Motala3(Male) 6,000BC Motala Sweden: Y DNA=I2a1b*(I M258+, I PF3742+, I2 L68+, I2a1 P37.2+, I2a1b CTS7218+, I2a1b CTS1293+, I2a1b CTS176+, I2a1b1 M359.2-, I2a1b3 L621-), mtDNA=U5a1

Motala4(Female) 6,000BC Motala Sweden: mtDNA=U5a2d

Motala6(Male) 6,000BC Motala Sweden: Y DNA=? (Q1 L232- Q1a2a L55+), mtDNA=U5a2d

Motala9(Male) 6,000BC Motala Sweden: Y DNA=I* (I P38+, I1 P40-), mtDNA=U5a2

Motala12(Male) 6,000BC Motala Sweden: Y DNA=pre-I2a1b or brother lineage to I2a1b(I PF3742+, I M258+, I M170+, I2 L68+, I2a L460+, I2a1 P37.2+, I2a1b CTS7218+, I2a1b CTS5985+. I2a1b L178+, I2a1b CTS1293+, I2a1b CTS176+, I2a1b CTS5375-, I2a1b CTS8486-, I2a1b1 M359.2-, I2a1b3 L621-), mtDNA=U2e1

Neolithic Swedish hunter gatherers of the Pitted Ware culture

Ajv52A(Male), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=V

Ajv59(Male), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U

Ajv53(Female), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U4d

Ajv58(Male), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: Y DNA=I2a1-P37.2, mtDNA=U4d

Ajv70(Male), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U4d

Ire8(Male), 5,100-4,150 cal. B.P, Ire, Hangvar, Gotland Sweden: mtDNA=U4d

Ajv13(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U4

Ajv52b(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U4

Ajv66(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U4

Ajv54(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U5

Ajv36(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U5

Ajv5(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U5a

Ajv29a(?), 4,900-4,600 cal B.P, Ajvide, Eksta, Gotland Sweden: mtDNA=U5a

Fir15(?), 2800-2000 BC,Fridtorp, Västerhejde, Gotland: mtDNA=U4

Fir22(?), 2800-2000 BC,Fridtorp, Västerhejde, Gotland: mtDNA=U4

Fir4(?), 2800-2000 BC,Fridtorp, Västerhejde, Gotland: mtDNA=U5

Fir27(?), 2800-2000 BC,Fridtorp, Västerhejde, Gotland: mtDNA=U5a

Ire6b(?), 5,100-4,150 cal. B.P, Ire, Hangvar, Gotland Sweden: mtDNA=T2b

Ire9(?), 5,100-4,150 cal. B.P, Ire, Hangvar, Gotland Sweden: mtDNA=U4

Ire3(?), 5,100-4,150 cal. B.P, Ire, Hangvar, Gotland Sweden: mtDNA=U4


Neolithic Swedish Farmers of the TRB culture, Frälsegården, Gokhem Sweden

Gökhem4(Male), 5,050-4,750 cal. years B.P.: mtDNA=H

Gökhem2(Female), 5,050-4,750 cal. years B.P.: mtDNA=H1c

Gökhem7(Female), 5,050-4,750 cal. years B.P.: mtDNA=H24

Gökhem5(Female), 5,280-4,890 cal. B.P.: mtDNA=K1e


Ste7(Female), 5,280-4,890 cal. B.P.: mtDNA=T2b

Ste7(Female), 5,280-4,890 cal. B.P.: mtDNA=J

The mtDNA of the Stone age Swedish farmers and hunter gatherers is 100% constant with what has been found in stone age farmers and hunter gatherers from other regions of Europe. The hunter gatherers have about 100% U5, U4, and U2, and the farmers mainly have modern European-specfic subclades of haplogroups which are most diverse and originated in the middle east. Since mDNA H1 and H3 are most popular in western Europe today, i bet that Neolithic west Europeans like the ones in Sweden had a very high amount and mtDNA samples from them so far is great evidence this is true. So far all U5 samples from Swedish hunter gatherers are U5a(mainly U5a1, but also U5a2 and U5a2d), like Mesolithic Russians(specifically U5a1) and unlike Mesolithic central-west Europeans who had mainly U5b(mainly U5b2) and the only U5a subclade found are U5a2(including U5a2c3* and U5a2a). The U5a in Swedish hunter gatherers may be from east European-very ANE like ancestors,, while their Y DNA I2a1-P37.2 is probably descended of central-west European ancestors.

At least paternally modern Balts and Scandinavians don't show evidence of a high amount of Mesolithic Swedish ancestry. I2a1-P37.2 today is very very rare in Scandnavians and Balts(much of it is probably east European=specfic I2a1b1a-L147.2), there are some Scandinavian P37.2 samples in FTDNA's database who have not been tested for I2a1-P37.2 subclades i bet at least have unknown specific Scandinavian SNPs and are a brotherclade to I2a1b. Y DNA I2a1-P37.2 is likely a marker of Europeans who took refuge in south-western Europe during the last ice age, today it is most diverse in western Europe and so far has been shown to be the main lineage of Mesolithic central-west-north Europeans.

Ancient and modern mtDNA are great evidence that Mesolithic Europeans at least maternally largely descended of humans who arrived in Europe over 30,000 years ago. There are no Y DNA samples from Upper Palaeolithic Europe, but the fact that Y DNA I is most diverse in Europe, 6/7 Mesolithic Europeans tested have Y DNA I, and that it is estimated to be 20,000-25,000 years old is great evidence that many Upper Palaeolithic Europeans belonged to Y DNA I. There are other ancient and exclusively European Y DNA haplogroups like C1a2-V20 which is what Mesolithic Spaniard La Brana-1 had, but also F-96 and maybe others i don't know of. Upper Palaeolithic Siberian MA1 belonged to Y DNA haplogroup R* and he was a pure west Eurasian, it is possibly that some Upper Paleolithic European belonged to Y DNA R, Q, or another descendant of P.

Autosomal DNA shows that Mesolithic Europeans all descended from the same ancient source, that their modern descendants are largely confined to Europe, and that they are likely descended of very early Europeans who arrived i think at least before the last Ice age.
Autosomal DNA
By using D-statistics Skoglund et al. 2014 found that Ajv58 fits as a clade with La brana-1, modern Sardinians fit as a clade with Otzi, and Gok2 has more hunter gatherer ancestry than Otzi. They found that all non-west Eurasian non Africans share more drift with Ajv58, which is because Gok2 had some basal Eurasian ancestry which does not have the shared drift between west and east Eurasians. The modern Europeans who share more dirft with either Ajv58 or Gok2 is constant with how Mesolithic European and near eastern ancestry is distributed in Europe. Northern and central Europeans are closer to Ajv58 while southern Europeans(incl. French) are closer to Gok2.

Not surprisingly modern Sardinians share much more dirft with Gok2 than any other modern populations. I was shocked to see that Sami(specifically from Norrbotten, Sweden) share much more drift with Ajv58 than any other modern Europeans, they share as much drift with Ajv58 as Sardinians do with Gok2. This suggests that Sami have the highest amount of Mesolithic European hunter gatherer ancestry today, and coincidentally are the last Europeans who are traditionally hunter gatherers.

Ajv58 may have had no farmer ancestry but Gok2 probably had around 30-40% hunter gatherer ancestry. So the D-statistics mean that Mesolithic European hunter gatherer ancestry is probably over 40% in much of northern and central Europe.

Skoglund et al. 2014 created admixture graphs like Lazaridis 2014 with ancient and modern samples, and came to the same conclusions.

Four key admixture events shown by their tree

1. Gene flow from Denisovans to Australian aboriginal populations (48, 97, 98)
2. Gene flow from the MA1 lineage to Native American ancestors (Anzick1 and
Saqqaq) (13)
3. Gene flow from the MA1 lineage to Ajvide58, see main text and Table S13.
4. Gene flow from Late Mesolithic hunter-gatherers (La Brana 1) to early
farmers (Gökhem2 and Iceman) (18).


Their admixture graphs like Lazardis suggest early European farmers harbored a significant amount of basal Eurasian ancestry(44+or-3% for Otzi, 23+or-6% for Gok2) and the rest of their ancestry formed a clad with European hunter gatherers. Most of their European hunter gatherer-like ancestry, descends from near easterns who were closely related to Mesolithic Europeans. Gok2 though is estimated to have 21% more than Otzi, and that 21% extra is defintley European hunter gatherer ancestry, Gok2 may have been as much as 40% Mesolithic European.

skoglund.png


In their fitted model Ajv58(not La Brana-1) is the best fit for the hunter gatherers that contributed ancestry to Gok2 and Otzi. This is because Gok2's and Otzi's hunter gatherer ancestors were probably from central Europe(for Gok2 also northern Europe and Scandinavia) and were more related to Ajv58 a Swedish hunter gatherer than to La Brana-1 an Iberian hunter gatherer. Similarly Lazardis found that Stuttgart is more related to Loschbour and Motala12 than to La Brana-1, because her hunter gatherer ancestors were also probably from around central Europe.

Skoglund et al. 2014 fitted Ajv58 a Neolithic Swedish hunter gatherer as a mixture of MA1 and La Brana-1, like how Lazaridis 2014 fit Mesolithic Swedish hunter gatherer Motala12 as a mixture of MA1 and Loschbour. The predicted percentages of ANE(~25%) and WHG(~75%) ancestry in Ajv58 are very similar to what Lazaridis 2014 predicted for Motala12(19% ANE, 81% WHG). It was already noticed by Laz that Neolithic Swedish hunter gatherers cluster very closely to Mesolithic ones because they shift more towards MA1 in PCAs than La Brana-1 and Loschbour do. La Brana-1 like Loschbour did not fit as having any ANE ancestry.

This PCA of the ancient genomes with modern west Eurasians, are constant with Lazardis's and Davidski's(Eurogenes blog) PCAs.
Skoglund_2014_PCA_small.png


24,000BP Siberian MA1 clusters more closely with European hunter gatherers than with near easterns, which is probably because he and European hunter gatherers(some farmer ancestry for La Brana-1, and probably for the Neolithic Swedish hunter gatherers) are pure west Eurasians, unlike middle eastern specific ancestry which has a high amount of basal Eurasian.

The Neolithic Swedish farmers are much farther up towards European hunter gatherers than Otzi is. They cluster most closely with Basque, who are one of few modern Europeans who fitted as a mix of Stuttgart and Loschbour in Laz, and have the highest amount of WHG ancestry in southern Europe. One of the Neolithic Swedish farmers is shifted about as far up as central-north Europeans. In my opinion the Swedish Neolithic farmers had 30%- 40% WHG ancestry. They may have more hunter gatherer ancestry than Otzi and Stuttgart because they were the first farmers in Scandinavia.

The D-statistic above shows that MA1 is much closer to Lithuanians(arguable the most Mesolithic-like modern Europeans) than to Sardinians(the most Neolithic-like modern Europeans), like European hunter gatherers. He and StoraForvar11 are almost exactly as distant from Sardinians but La Brana-1 and the Swedish Neolithic hunter gatherers are less distant from Sardinians, probably because they had some farmer ancestry. MA1 has constantly been shown to be a close relative of stone age European hunter gatherers, but this is because he and stone age European hunter gatherers are the only samples known of pure or close to pure west Eurasians. Middle eastern-specific ancestry comes primarily from a source more related to European hunter gatherers than MA1 is, but its basal Eurasian ancestry makes them less related to European hunter gatherers than MA1 is.

Pigmentation

Ajv58 had ancestral "dark skin" alleles in SNPs rs1426654, like La Brana-1 and Loschbour but unlike Mesolithic Swede Motala12 who had the derived "light skin" alleles. Ajv58 also had ancestral 'dark skin" alleles in SNP rs16891982 like Loschbour and La Brana-1 but unlike Mesolithic Swede Sf11 who had the derived "light skin" alleles. So most Mesolithic Europeans had the ancestral "dark skin" alleles in those two SNPs, but a minority had the derived "light skin" alleles.

Early European farmers Otzi, Gok2, and Stuttgart all had the derived "light skin" alleles in SNP rs1426654, Stuttgart had the ancestral 'dark skin" alleles in SNP rs16891982, while Gok2 and Otzi had the derived light skin alleles in SNP rs16891982. "Light skin" alleles in SNP rs1426654 were probably fixated in early European farmers, as they are in modern Europeans and west Asians. SNP rs16891982 is also associated with hair color, one study claims that if a European individual has the ancestral alleles there is a 7x better chance that individual has black hair. That's why the derived alleles are less popular in southern Europe than in northern Europe, and less popular in the middle east than in Europe.

Maybe since these two "light skin" mutations existed in both European hunter gatherers and farmers but at differnt rates, when the two populations mixed their descendants had these mutations at the same rate of their farmer ancestors. Like i have said in this thread and other times the science behind human skin color is not very well known and it's impossible to say what skin color these ancient people had.

I think i may have relatives who are good proxies for the skin color of most Mesolithic Europeans had. My brown skinned uncle is 92% European, 8% Native American+African, and he is missing the Ala111Thr and Phe374Leu mutations like La Brana-1, Loschbour, and Ajv58, and he is missing other mutations associated with European light skin and are fixated in modern Europeans. He is probably one of few people who have a significant amount of European hunter gatherer ancestry(probably around 40% or more) and is missing many of the same light skin mutations.

Neolithic European farmers defintley had light skin, since they had all the mutations associated with light skin in west Eurasia today and modern Sardinians are pracituclley no differnt genetically are generally light skinned. I tend towards saying stone European hunter gatherers had dark skin, because it cant be random that they are missing nearly every mutations associated with European light skin that are fixated in modern Europeans. There are alot of possibilities though, maybe both hunter gatherers and farmers had light skin. Today it seems all light pigmentation in Europe correlates with WHG ancestry, so possibly the hunter gatherers had light skin and some light hair, not just light eyes.

I was hoping Skoglund et al. 2014 would help find the origin of high amounts of light hair in modern Europeans. In supplementary pages 36-37 they have a horrible pigmentation predictor, so that's disappointing. In my opinion light hair in Europe today is either mainly descended of European hunter gatherers, or first became popular in hunter/farmer mutts. Since 3/4 Stone age European hunter gatherers tested so far are missing the L374F they were probably very dark haired. Near eastern ancestry today in Europe correlates with dark hair, Sardinians are the most early European farmer descended modern Europeans and are also the darkest haired Europeans so early European farmers were also probably very dark haired.
 
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Seems as if the (PWC/Hunter-gatherer p.19) Ajvide58 is lacking the light-skin allele rs1426654/SLC24A5 being G/G like Loschbour and LaBrana - a contrast to Motola12 (A/A) despite being ~1000 years before Ajv58; Whereas Gök2 (TRB/Neolithic-farmer p.18) has the light-skin rs1426654/SLC24A5 (A/A) likewise as Ötzi and Stuttgart while Gök4 and the others dont have a result;

I would not call it as great as Lazaridis 2013 most of Skoglund 2014 it is consistent with all else and expected after Lazaridis and Rhagavan;
 
Great stuff, lots of new information, seemingly nothing inconsistent with what we've already been learning. Only one Y-DNA result this time: I2a1, which quite possibly could have been I2a1b* again if they had gotten more SNP results. It could also have been I2a1a, though, which is more typically thought to have been absorbed by, and spread with, the farmers.

Pitted Ware is interesting because they were hunter-gatherers living at a time when the Neolithic had spread right up next to them.
 
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Good stuff, quite in line with other results as pointed above. The most revolutionary finding still seems to be La Braña's paternal line though, which has no equal until date.

Haplogroup I variants are showing strong presence during the Mesolithic in Northern Europe. One can speculate if C-V20 was restricted to Southern Europe, or if a population replacement already put haplogroup I in the scene regarding the new samples. So it would be interesting to test other Mesolithic individuals from both Northern and Southern Europe, and see if before 6000 BC, C-V20 shows up pretty much dominant, specially in Northern Europe. If that eventually becomes confirmed, it would indicate that the population replacement started from the North and then gradually to the South. There's also the possibility that haplogroup C-V20 is so incredibly old that already was a rarity during the Mesolithic, making very difficult to know when exactly its number decreased drastically.

Only one Y-DNA result this time: I2a1, which quite possibly could have been I2a1b* again if they had gotten more SNP results. It could also have been I2a1a, though, which is more typically thought to have been absorbed by, and spread with, the farmers.
I missed your interventions. I'd like to see that as well.
 
Good stuff, quite in line with other results as pointed above. The most revolutionary finding still seems to be La Braña's paternal line though, which has no equal until date.

Haplogroup I variants are showing strong presence during the Mesolithic in Northern Europe. One can speculate if C-V20 was restricted to Southern Europe, or if a population replacement already put haplogroup I in the scene regarding the new samples. So it would be interesting to test other Mesolithic individuals from both Northern and Southern Europe, and see if before 6000 BC, C-V20 shows up pretty much dominant, specially in Northern Europe. If that eventually becomes confirmed, it would indicate that the population replacement started from the North and then gradually to the South. There's also the possibility that haplogroup C-V20 is so incredibly old that already was a rarity during the Mesolithic, making very difficult to know when exactly its number decreased drastically.

There was some Y DNA diversity in Mesolithic Europe, but autosomally they didn't change much from Spain-Sweden. La Brana-1 is just one sample i would not be surprised if La Brana-2 will turn out to have your haplogroup I2a1a1-M26. There are many I2 subclades in central-western Europe, i am sure most of them have been in western Europe since the Mesolithic. I have heard the Cheddar man(Britian, 8,000BC) is going to be retested, if they get his Y DNA haplogroup i bet it will be sometype of I2a1, probably I2a1c-L1294 because it's the main-type of I2a1 in the British isles today.
 
One revelation from that study is that Gök4 who was previously (Skoglund 2012) considered a woman is actually a man (p.18); Gök2 app. remains a female;
 
One revelation from that study is that Gök4 who was previously considered a woman is actually a man (p.18);

The reverse is true for Micheal Jackson.

I think you'd be interested to learn my brown skinned uncle is missing the Ala111Thr and Phe374Leu mutations like La Brana-1, Loschbour, and Ajv54 and unlike nearly all modern Europeans. He is also missing many other light skin mutations, some also fixated in modern Europeans. I suspect that he has a similar skin color as did Mesolithic Europeans and may be one of few(not literally, but you get what i'm saying) remaining Europeans with this trait.
 
As with la Brana 1 I was surprised to find that a number of disease resistance genes have been found among hunter-gatherers. I somehow, in my imagination, have always compared the influx of farmers in Europe with the colonisation of North-America, where Indians are the WHGs and the whites are the EEFs. This analogy obviously works only on a higher level. But the difference in outcome is what is so surprising. Where American Indians - when defined as broad as possible - only make up 1,5% of the North-American population WHGs actually roughly make up one third of Europe's genetic material.

That is a colossally successful outcome of if you compare that to other hunter-gatherers around the world. The Indians were largely exterminated by the smallpox. One may be tempted to point that to their isolation, but a similar event happened to Siberian native tribes that are closely related to Indian. And similar things happened (and even happen today) to almost all tribes that cane into first contact with farmers. Small pox and even diseases such as common cold decimated (and continue to decimate) the tribes of the Andamans, the San and Bushmen, the last Indian tribes of the Amazones, the Negrito's, the Aboriginals and countless others.

Jared Diamond used this to partly explain the cultural dominance of the European civilization in his slightly flawed but highly readable book Guns Germs and Steel.

So, now we see hunter-gatherers that actually blended in. And consider this. The remaining American Indians are largely admixed with Europeans whereas present day American population is hardly admixed with Indians (a situation, mind you, that is quite different in Latin America). But after centuries contact WHG were still rather a pure breed and it was actually the farmers that mixed.

The European hunter-gatherers were a hugely successful breed, and the disease resistance genes play an important role in that success, I think
 
There was some Y DNA diversity in Mesolithic Europe, but autosomally they didn't change much from Spain-Sweden. La Brana-1 is just one sample i would not be surprised if La Brana-2 will turn out to have your haplogroup I2a1a1-M26. There are many I2 subclades in central-western Europe, i am sure most of them have been in western Europe since the Mesolithic. I have heard the Cheddar man(Britian, 8,000BC) is going to be retested, if they get his Y DNA haplogroup i bet it will be sometype of I2a1, probably I2a1c-L1294 because it's the main-type of I2a1 in the British isles today.

Where did you get that?
Cheddar man would be interesting. Cheddar 10000 years ago. My bet would be I2a1b , Magelmosian culture , but neither L161.1 nor L621 , some extinct branch instead.
Both L161.1 and L621 split only 6000 years ago and those don't seem to have happened on the Brittish Isles, they immigrated later.
I2a1c-L1294 should be reclassified. Both I2a1a1-M26 and L1294 seem to share the I2a1-CTS595 mutation.
It would be interesting to know where and when CTS595 split : was only M26 connected to the neolithic or was it the whole CTS595 branch?
 
There was some Y DNA diversity in Mesolithic Europe, but autosomally they didn't change much from Spain-Sweden.

They all came from Southern France and Northern Spain and moved after the younger dryas ( 12.700 -11.600 year ago ) and even some 1000 years later, when trees started to grow again up north.
They had lived more than 10000 years in this small area of Southern France and Northern Spain, with lots of intermarriages.

The first wave up north from Southern France were hunting reindeer on the tundra 15500 year ago , but they got extinct after the second wave came with the first trees.
Few of them survived tough in the Norvegian fjords. They resurged 4000 years ago. They were I1. They were fishermen and sailors by then.
 
Ancient and modern mtDNA are great evidence that Mesolithic Europeans at least maternally largely descended of humans who arrived in Europe over 30,000 years ago. There are no Y DNA samples from Upper Palaeolithic Europe, but the fact that Y DNA I is most diverse in Europe, 6/7 Mesolithic Europeans tested have Y DNA I, and that it is estimated to be 20,000-25,000 years old is great evidence that many Upper Palaeolithic Europeans belonged to Y DNA I. There are other ancient and exclusively European Y DNA haplogroups like C1a2-V20 which is what Mesolithic Spaniard La Brana-1 had, but also F-96 and maybe others i don't know of. Upper Palaeolithic Siberian MA1 belonged to Y DNA haplogroup R* and he was a pure west Eurasian, it is possibly that some Upper Paleolithic European belonged to Y DNA R, Q, or another descendant of P.

What mesolithic F-96 is there? I know of F-96 LBK neolithic.
 
Mind you, the notion that North-Europeans have one third to two fifths of WHG genetic material brings a number of problems with it. This study specifically claims that farmers absorbed WHGs as they spread rather than the other way around. But that still leaves one of the problems that arises.

A while ago another study, I think it was also by Skoglund, claimed that the genetic investigation of Gotland Pitted Ware Culture showed that at the onset of the neolithic the population was completely replaced. [1] They showed tables of the rate of mtDNA in current day as compared to the baltics and the PWC finds to prove that point. However, when la Brana was published however, it clearly showed close relation to Northern Europeans rather than Iberians. How can both those assumptions be true?

I have been wondering about this a lot. Where has the mtDNA gone? Swedes have rather a high WHG rate, yet very low U5 (or U4) mtDNA. If Skoglund is right and farmers mobbed up WHGs we would expect rather a high number of U5/4.

I notices the Saami, who may autosomically not be a good proxy for WHG but clearly are decended from them, have 50% U5 but also 50% V. The latter is considered to be introduced by farmers, but the high incidence of V among the Saami is considered to be the result of a founder effect. This led me to thinks about the Ertebolla culture and the Swifterband culture, two cultures that existed alongside the LBK culture but archeological evidence clearly shows continuation and slow adaptation to farming. They kept pigs and cultivated barley rather than wheat. What if especially the Ertebolla culture became so successful that LBK and/or Funnel Beaker women married into the culture, and due to the same founder effect their mtDNA became the main mtDNA? We do know from DNA from pig remains that at first domesticated pigs - both from LBK and Ertebolla - showed great affinity to the Near-east while later local DNA prevails in both cultures [2]. This at least shows that both cultures had a lot of contact.

So, I think the lack of U5 in Sweden may be due to the success of local hunter-gatherers in adapting rather than their replacement.


[1] http://dienekes.blogspot.nl/2012/04/ancient-dna-from-neolithic-sweden.html
[2] http://geknitics.com/2007/09/ancient-pig-dna-and-the-neolithic-transition/
 
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Mind you, the notion that North-Europeans have one third to two fifths of WHG genetic material brings a number of problems with it. This study specifically claims that farmers absorbed WHGs as they spread rather than the other way around. But that still leaves one of the problems that arises.

A while ago another study, I think it was also by Skoglund, claimed that the genetic investigation of Gotland Pitted Ware Culture showed that at the onset of the neolithic the population was completely replaced. [1] They showed tables of the rate of mtDNA in current day as compared to the baltics and the PWC finds to prove that point. However, when la Brana was published however, it clearly showed close relation to Northern Europeans rather than Iberians. How can both those assumptions be true?

I have been wondering about this a lot. Where has the mtDNA gone? Swedes have rather a high WHG rate, yet very low U5 (or U4) mtDNA. If Skoglund is right and farmers mobbed up WHGs we would expect rather a high number of U5/4.

I notices the Saami, who may autosomically not be a good proxy for WHG but clearly are decended from them, have 50% U5 but also 50% V. The latter is considered to be introduced by farmers, but the high incidence of V is considered to be the result of a founder effect. This led me to thinks about the Ertebolla culture and the Swifterband culture, two cultures that existed alongside the LBK culture but archeological evidence clearly shows continuation and slow adaptation to farming. They kept pigs and cultivated barley rather than wheat. What if especially the Ertebolla culture became so successful that LBK women married into the culture, and due to the same founder effect theit mtDNA became the main mtDNA? We do know from DNA from pig remains that at first domesticated pigs - both from LBK and Ertebolla - showed great affinity to the Near-east while later local DNA prevails in both cultures [1]. This at least shows that both cultures had a lot of contact.

So, I think the lack of U5 in Sweden may be due to the success of local hunter-gatherers in adapting rather than their replacement.


[1] http://dienekes.blogspot.nl/2012/04/ancient-dna-from-neolithic-sweden.html
[2] http://geknitics.com/2007/09/ancient-pig-dna-and-the-neolithic-transition/

Swifterbant and Ertebolle exosted after Doggerland had drowned, that is were they came from and i suspect I2a2a M223 descendants.
Swifterbant had trading contacts with neolithics and adopted cattle breeding and some wheat cultivation but kept their own identity.
It seems to me they expanded after the neolithics with IE people. The same goes for the I1 fishermen from the Norvegian fjords.

Saami only arrived 2600 year ago, coming from the east. U5 and V may have been there before together with a male population now extinct.
 
The greatest variety of I1 seems to exist in Northern Germany or thereabouts. If the origin of its spread was in Western Norway, one would expect to find diversity or basal subclades in greater numbers there.
 
Swifterbant and Ertebolle exosted after Doggerland had drowned, that is were they came from and i suspect I2a2a M223 descendants.
Swifterbant had trading contacts with neolithics and adopted cattle breeding and some wheat cultivation but kept their own identity.

The DNA from pigs found at an Ertebolla site showed it to be of Near-Eastern decent, which points to trading as well. Mind you, Swifterband excavations actually yielded bones so I have the hope that someone tries to extract DNA from them.

It seems to me they expanded after the neolithics with IE people. The same goes for the I1 fishermen from the Norvegian fjords.

Saami only arrived 2600 year ago, coming from the east. U5 and V may have been there before together with a male population now extinct.

Saami are considered a mixture of a Siberian component and a local component, from what I understand.
 
The greatest variety of I1 seems to exist in Northern Germany or thereabouts. If the origin of its spread was in Western Norway, one would expect to find diversity or basal subclades in greater numbers there.

Todays I1 all descend from only 1 tribe some 4000 year ago. A highly mobile tribe sailing the Baltic. It spread very fast around the Baltic and later also expanded south.

For their origin look at Ahrensburg and Fosna-Hensbacka culture. Ahrensburg people dissapeared when the forests started to grow north. Other people replaced them.
 
The greatest variety of I1 seems to exist in Northern Germany or thereabouts. If the origin of its spread was in Western Norway, one would expect to find diversity or basal subclades in greater numbers there.

I know trying to figure out where I1 originated is very annoying. It was obviously spread by Germanic people(not counting Finnish-specific subclades and other rarer subclades) but is most diverse in central Europe, and Germanic people spread from southern Scandinavia and far northern central Europe. Y DNA of Swedish hunter gatherers have proven I1 probably came to Scandinavia in the metal ages.
 
As with la Brana 1 I was surprised to find that a number of disease resistance genes have been found among hunter-gatherers. I somehow, in my imagination, have always compared the influx of farmers in Europe with the colonisation of North-America, where Indians are the WHGs and the whites are the EEFs. This analogy obviously works only on a higher level. But the difference in outcome is what is so surprising. Where American Indians - when defined as broad as possible - only make up 1,5% of the North-American population WHGs actually roughly make up one third of Europe's genetic material.

That is a colossally successful outcome of if you compare that to other hunter-gatherers around the world. The Indians were largely exterminated by the smallpox. One may be tempted to point that to their isolation, but a similar event happened to Siberian native tribes that are closely related to Indian. And similar things happened (and even happen today) to almost all tribes that cane into first contact with farmers. Small pox and even diseases such as common cold decimated (and continue to decimate) the tribes of the Andamans, the San and Bushmen, the last Indian tribes of the Amazones, the Negrito's, the Aboriginals and countless others.

Jared Diamond used this to partly explain the cultural dominance of the European civilization in his slightly flawed but highly readable book Guns Germs and Steel.

So, now we see hunter-gatherers that actually blended in. And consider this. The remaining American Indians are largely admixed with Europeans whereas present day American population is hardly admixed with Indians (a situation, mind you, that is quite different in Latin America). But after centuries contact WHG were still rather a pure breed and it was actually the farmers that mixed.

The European hunter-gatherers were a hugely successful breed, and the disease resistance genes play an important role in that success, I think

I think there's something to this in terms of the effect of the disease resistant genes allowing them to retain a genetic presence in Europe. (I have to check.Did the earliest sample have them? I think that might be important, because the 6,000 B.C.samples already had one T2b mtDNA among them. I've longthought that the Native Americans who survived did so because they had some proportion of European genes. That's true in Latin America as well, where many tribes previously considered to be 100% Native American have been found to harbor some European genes. Of course, in some areas of Latin America, many of the native had already discovered agriculture, leading to larger population numbers, and so were not as easily wiped out.

However, what is striking to me is that this all seems to show that after decades of research, new technology, and more sophisticated mathematical modeling, I seem to be back to where I started with all of this, with Cavalli Sforza. Isn't this all a validation of the demic diffusion model of the advance of agriculture? This is what the authors have to say...


"This suggests that Gökhem2 has additional hunter gatherer ancestry which is not present in the Iceman...


Finally, we tested for each modern day population in the SNP array data set whether it shared more genetic drift with Gökhem2 or Ajvide58. We find that
Swedish populations generally tend to share most genetic drift with Ajvide58 Table S14). In contrast, many central and southern European
populations do not show specific affinity to either group using this test, whereas populations in the Middle East, Turkey, Cyprus, and Armenia
show stronger affinity to Gökhem2. This suggests that the initial pulse of ancestry that is associated with early farmers was further diluted
in descendant populations in Northern Europe."


The higher numbers for WHG as you move north in Europe fit rather nicely within that framework, I think.

There's also the fact that because of the unsuitability of northern Europe for the initial Neolithic agricultural package, these WHG, minority ANE people were left alone for a very long time. It wasn't until crops were developed that would thrive in that envrionment that parts of Europe were neolithicized, and it was only with that changeover that their numbers increased.

It's easier to retain your genetic cohesiveness when no one wants the land you're sitting on, or at least is at the limits of what is desired.
Even in central Europe,there was a climate collapse, and perhaps an environmental crisis created deforestation. All of these factors come into play it seems to me, as well as the fact that there was a later movement of peoples into Europe who were not part of the original encounter at all.

I think it also has to be borne in mind that we are talking abouti yDNA lineages that are very much minority lineages in Europe today. The "R"lineages have nothing to do with any of this. That change over happened later, and with an intrusive group, or at least that seems to be the case from the evidence we have to date.

Although I think we should be cautious about making broad generalizations, it's interesting that while you had a "farmer" mtDNA in the hunter-gatherers, you didn't yet have the hunter-gatherer mtDNA in the farmers. The same thing happened in the Balkans according to a prior study. In the earliest stages of the encounters, the mtDNA gene flow seemed to go only in one direction, although of course that changed later. Is it possible that it was the "farmer" women who brought agriculture to the natives? Of course, in the Balkans, they did also incorporate some I2a yDNA lineages. I wonder if that was because, given the climate in the Balkans, the agricultural package was even more obviously successful there.
 
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I know trying to figure out where I1 originated is very annoying. It was obviously spread by Germanic people(not counting Finnish-specific subclades and other rarer subclades) but is most diverse in central Europe, and Germanic people spread from southern Scandinavia and far northern central Europe. Y DNA of Swedish hunter gatherers have proven I1 probably came to Scandinavia in the metal ages.

I1 L22+ Z74+ is ancestral to both the main Finnish subclade (L287+) and one of the large Scandinavian subclades (L813+). The questions there are when the split happened and whether Z74+'s spoke Indo-European or something paleo-European before the split.

L300 (which is Z74- like P109 and L205) is IMO not important when it comes to the bigger story of I1 in the Baltic region. There's only a few individuals with it in southern Finland, and one who traces his ancestry into Sweden. I think it probably came to Finland with Swedes less than 1000 years ago and only shows up in greater numbers here because Sweden is undersampled in comparison to Finland in FTDNA projects.
 

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