Fire Haired
01-26-2014, 08:43 AM
Lazaridis 2013's abstract.
Analysis of ancient DNA can reveal historical events that are difficult to discern through study of present-day individuals. To investigate European population history around the time of the agricultural transition, we sequenced complete genomes from a ~7,500 year old early farmer from the Linearbandkeramik (LBK) culture from Stuttgart in Germany and an ~8,000 year old hunter-gatherer from the Loschbour rock shelter in Luxembourg. We also generated data from seven ~8,000 year old hunter-gatherers from Motala in Sweden. We compared these genomes and published ancient DNA to new data from 2,196 samples from 185 diverse populations to show that at least three ancestral groups contributed to present-day Europeans. The first are Ancient North Eurasians (ANE), who are more closely related to Upper Paleolithic Siberians than to any present-day population. The second are West European Hunter-Gatherers (WHG), related to the Loschbour individual, who contributed to all Europeans but not to Near Easterners. The third are Early European Farmers (EEF), related to the Stuttgart individual, who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model the deep relationships of these populations and show that about ~44% of the ancestry of EEF derived from a basal Eurasian lineage that split prior to the separation of other non-Africans.
If you want to download Lazaridis 2013 click here (http://biorxiv.org/content/early/2013/12/23/001552).
The first thread I made about Laz 2013(Y DNA I found in Mesolithic Sweden(I1-) and Luxemburg, a lot of other good stuff (http://www.theapricity.com/forum/showthread.php?107551-Y-DNA-I-found-in-Mesolithic-Sweden(-I1)-and-Luxemburg-alot-of-other-good-stuff)) was terrible because I posted it less than a day after Laz 2013 was published and I couldn't actually download the study so I got all of my information from posters. Even though now I have Laz 2013 downloaded I could study it for days, weeks, months, even years and still have many questions and unconfident conclusions. The most confusing thing to me are the ghost populations ANE and basal Eurasian. There are so many concepts involved with Laz 2013 that I can't remember it all in my head I have to write it down. The paper on La Brana-1's genome is suppose to be out tomorrow January 27th, 2014. I am sure the researchers compared La Brana-1's genome to data in Laz 2013, so it will help learn even more about European and overall human genetic history. It has been said by researcher Carles Lalueza-Fox La Brana-1 had the blue eye mutation(I assume it is the mutation in SNP rs12913832 (http://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=4&cad=rja&ved=0CEgQFjAD&url=http%3A%2F%2Fsnpedia.com%2Findex.php%2FRs12913 832&ei=QtjkUrWbJ8bhsAS4toKgCg&usg=AFQjCNHtVzdx9_RYDSyYm4KR8CKSXGWxkg&sig2=-fZWTGABJhLx4HQ9a-a4Ng&bvm=bv.59930103,d.aWM)) and Eupedia member CLF who claims to be Carles Lalueza-Fox said La Brana-1 had Y DNA C-V20 and that the paper should be out tomorrow.
Below are the three ancestral populations Laz 2013 claims modern Europeans descend from and the ancient individuals their based on, I coped and pasted this list from this (http://eurogenes.blogspot.com/2014/01/another-look-at-lazaridis-et-al-ancient.html) Eurogenes thread. The list includes ENA because north-east Europeans were shown to have some ENA ancestry(had already been discovered) also Sicilians, Maltese, and Ashkenazi Jews were found to maybe have some near eastern ancestry that can't only be explained by EEF.
- West European Hunter-Gatherer (WHG), based on an 8,000 year-old genome from Loschbour, Luxembourg
- Ancient North Eurasian (ANE), based on a 24,000 year-old genome from South Siberia (dubbed Mal'ta boy or MA-1)
- Early European Farmer (EEF), based on a 7,500 year-old genome from Stuttgart, Germany, belonging to the Neolithic Linearbandkeramik (LBK) culture
- Eastern non-African (ENA), this basically means East Eurasian, and is based on samples of present-day Onge, Han Chinese and Atayal from Taiwan
I have been confused who MA1 was genetically since the study that originally sampled his genome Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) was published in November 2013. In admixtures he looks like a mixture of European(specifically components that are most popular in north-eastern Europe and probably mainly of Mesolithic European origin), south-central Asian, native American, and maybe a little Oceania. I have been told by Davidski at Euorgenes that he is not a mix of those populations it is just that those components have MA1 like admixture.
Here are some important things I learned from Laz 2013 about the relationship between the ancient samples MA1, Loschbour, Motola12, and Stuttgart with eastern non Africans.
>Eurasian hunter gatherers(MA1, Stuttgart, and Motola12) are equally related to eastern non Africans,
>Eastern non Africans are more related to the Eurasian hunter gatherers(MA1, Loschbour, and Motola12) than to Stuttgart
>Eurasian hunter gatherers(MA1, Loschbour, and Motola12) and Stuttgart are more related to native Americans than they are to other eastern non Africans
>Native Americans are more related to MA1 than to European hunter gatherers(Loschbour and Motola12) and Stuttgart
>Papuans are less related to the Eurasian hunter gatherers(MA1, Loschbour, and Motola12) and Stuttgart than other eastern non Africans are Laz 2013 suggests that Denisovans admixture is the reason for this
>Atayal might be slightly less related to the ancient Eurasians(MA1, Loschbour, Motola12, and Stuttgart) than other eastern non Africans are.
Here is a figure from Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) "showing the levels of shared genetic drift between MA-1 and 147 present-day non-African populations", quote from Eurogenes thread First genome of an Upper Palaeolithic human. (http://eurogenes.blogspot.com/2013_11_01_archive.html)
http://img811.imageshack.us/img811/4151/zubo.png
MA1 is most related to native Americans(don't exactly know what shared drift means) but had no ENA ancestry. This means native Americans have some non ENA ancestry that is related to MA1. MA1 had mtDNA U(apart of his own distinct subclade) which is a west Eurasian mtDNA haplogroup(assumed, because it is so restricted to west Eurasians and most diverse in west Eurasians) he also had Y DNA R(R1-, R2-) which has a very debated origin, I don't think it originated in west Eurasians R more likely originated in a eastern non African people in my opinion because it descends from K-M526 and K-M256's except R and maybe Q(so their common ancestor P) originated in eastern non Africans. MA1 is 24,000 years old and R its self is estimated to be around 30,000 years old so maybe it originated in a people related to MA1 so maybe very related to west Eurasians and not eastern non African at all.
I don't know if MA1 was a west Eurasian person, from an extinct branch of the Eurasian family, or from a brother branch to west Eurasians. Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) says in their abstract there is evidence in MA1's autosomal DNA that he was basal to west Eurasians, closely related to native Americans, and had no affinity to east Asians. In PCA's MA1 and a 17,000 year old Siberian named AG2 clustered somewhat close to west Eurasians. Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) also said in their abstract that AG2 had similar autosomal genetic signatures as MA1. So I think MA1 was probably apart of a large population that was most related to west Eurasians and lived in Siberia(maybe other places) at least from 24,000-17,00 years ago and has left ancestry in many modern Eurasians and also in native Americans. I have noticed not many people consider that MA1 was a mutt. I think he may have had ancestry from many different(but related) populations that contributed ancestry to modern people separately.
Laz 2013 and Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) show that the relation to MA1 varies in west Eurasians which I think is prove there is also MA1 related ancestry in some west Eurasians. Relatedness to MA1 in the ancient European samples also varied because Stuttgart and Loschbour had little to no MA1 related ancestry but MA1 shared more alleles with Motola12 than he did with Loschbour. It was estimated that Motola12 had 81% Loschbour related ancestry and 19% MA1 related ancestry.
Laz 2013 found in many different tests that all modern Europeans(except maybe Basque, French_South, Bergamo, Pais_Vasco, Sardinian) are not an admixture of just Loschbour and Stuttgart related populations but also have MA1 related ancestry. Some Europeans scored a lower F3-statistic with Stuttgart-MA1 than Stuttgart-Loschbour showing some Europeans have more relation to MA1 than Loschbour and evidence for MA1 related ancestry. The F4-statistic(test population, Stuttgart, MA1, Chimp) is positive in many Europeans and near easterns but when MA1 is replaced by Loschbour it is only positive in Europeans. Meaning Loschbour related ancestry may be restricted to Europe but MA1 and Stuttgart related ancestry exists in Europe and the near east.
In F-3 statistics that included every possible modern west Eurasian population with at least four samples and one of the ancient samples as a at least one of the references, every near eastern population had its lowest statistic with Stuttgart as one of the references none had Loschbour or Motola12 as one of the references. In a F4 statistic it was shown that most modern Europeans have more Loschbour related ancestry than Stuttgart does but it also showed all modern near easterns have less Loschbour related ancestry than Stuttgart had. Evidence there is no close relationship between Loschbour and modern near easterns, Stuttgart derived from mainly the same source that modern near easterns derive from, and that Stuttgart had Loschbour related ancestry it was predicted that she had less than 100% near eastern ancestry and possibly as little as 61%.
I am very surprised by how unrelated modern near easterns are to Loschbour because in many admixtures like globe13 the component(s) that are most popular in north-eastern Europe and are mainly Mesolithic Europeans descended are the closest relative(s) to the component(s) that are most popular in northern west Asia. Since the similar results came from so many different admixtures I thought there must be some type of close relationship between Mesolithic Europeans and northern west Asians.
I am also very surprised by how related Stuttgart is to modern near easterns and that near easterns all probably descend from the same source. This is exactly what many people assumed though because farming spread to Europe from the near east. mtDNA and Y DNA of Neolithic European farmers does show that they were more related paternally and maternally to modern near easterns than any modern near easterns are to Mesolithic Europeans. It would make more sense to me that the near east is very diverse with many different west Eurasian family groups but Laz 2013 gives evidence they all come from the same root and that European(Loschbour) is their brother clade.
I really don't understand why Stuttgart would have ~44% basal Eurasian ancestry, from a people that separated from the Eurasian root before the east and west split. If this is true that means there is a large amount of basal Eurasian ancestry in Europeans, near easterns, and probably north Africans. I guess basal Eurasian ancestry could be an explanation of why Stuttgart is less related to eastern non Africans than ancient Eurasian hunter gatherers(MA1, Loschbour, and Motola12) were. I haven't seen anyone question how basal Eurasian ancestry would affect looks. I doubt Sardinians and Stuttgart would have looked like typical west Eurasians(which they do) with having ~44% non west Eurasian ancestry. If the basal Eurasian thing is somewhat true then I think possibly Y DNA E1b1b and G2a and mtDNA N1a1a and W in Neolithic Europeans may be connected.
No First I will say for native Americans there is no EEF or WHG ancestry just ANE which was predicted by Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) to be about 14-38% of their total ancestry.
Laz 2013 made an admixture test EEF(Stuttgart), WHG(Loschbour), and ANE(MA1) it really only works for Europeans though and for some Europeans who have extra near eastern ancestry or east Asian ancestry the results will be inaccurate. Eurogenes project allows members who have Eurogenes K13 results(click here (http://bga101.blogspot.com/2013/12/eef-whg-ane-test-for-europeans.html)) to get results in this test.
Iberia and France
<tbody>
Population
EEF
WHG
ANE
Basque
59.3
29.3
11.4
Pais_Vasco
71.3
12.5
16.3
Spanish
80.9
6.8
12.3
French
55.4
31.1
13.5
French_South
67.5
19.5
13
</tbody>
North west Europe
<tbody>
Population
EEF
WHG
ANE
Icelandic
39.4
45.6
15
English
49.5
36.4
14.1
Norwegian
41.1
42.8
16.1
Orcadian
45.7
38.5
15.8
Scottish
39
42.8
18.2
</tbody>
North east Europe
<tbody>
Population
EEF
WHG
ANE
Lithuanian
36.4
46.4
17.2
Estonian
32.2
49.5
18.3
</tbody>
Slavic east Europe(X Balkans)
<tbody>
Population
EEF
WHG
ANE
Ukrainian
46.2
38.7
15.1
Belorussian
41.8
43.1
15.1
Czech
49.5
33.8
16.7
</tbody>
Balkans an Italy
<tbody>
Population
EEF
WHG
ANE
Bergamo
71.5
17.7
10.8
Bulgarian
71.2
14.7
14.1
Croatian
56.1
29.3
14.5
Greek
79.2
5.8
15.1
Tuscan
74.6
13.6
11.8
Sardinian
81.7
17.5
0.8
Sicilian
90.3
0
9.7
Albanian
78.1
9.2
12.7
</tbody>
Near east
<tbody>
Near east
EEF
WHG
ANE
Ashkenazi_Jew
93.1
0
6.9
Maltese
93.2
0
6.8
</tbody>
Laz 2013 like many other people have found that Loschbour(or European hunter gatherers in general) are most related to modern northern Europeans especially north eastern and Scandinavian and that Stuttgart(or Neolithic European farmers in general) are more related to modern southern Europeans especially southwestern like Basque and Sardinia. In the PCA's Sardinians cluster very closely to Stuttgart and in K=2-K=20 admixtures from Laz 2013 had nearly identical results like they did with Otzi(~5,300BP early copper age farmer from the alps) and ~5,000 year old Swedish Funnel beaker farmers. I think Sardinians are nearly completely un mixed descendants of Stuttgart like farmers and during the Neolithic all central-west-northern(maybe other regions) European farmers were Stuttgart/Sardinia like.
The distribution of WHG ancestry is nearly identical to component(s) in many different admixtures usually called north European or another name referring to an area in northern Europe. The modern componets though probably have some non European hunter gatherer admixture. When near easterns are put in this admixture EEF, WHG, and ANE they usually have over 100% EEF and negative WHG this is constant with other evidence that seem to say there is little to no WHG ancestry in the near east.
ANE unlike WHG and EEF has a pretty even distribution in Europe. It seems to be a little bit higher in northern Europe though especially in north-east Europeans. ANE ancestry though is probably by far highest in native Americans and is also probably higher in the Caucasus than anywhere in Europe. When put in an admixture with two options near east and ANE Caucasus populations Abkhasian scored 18.6% ANE, Chechen scored 27%, and Lezgin scored 28.8%. There is also scattered ANE ancestry in the near east but at a lower levels than in what most Europeans have.
This is a map of the figure from Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) from the begging of this thread which shows that MA1 is more related to Europeans than near easterns, and there is a PCA from Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013).
http://3.bp.blogspot.com/-qdDK2ZHSeV4/UpxqAm3iUvI/AAAAAAAACRY/pthyx4u0VzU/s1600/Mal%27ta1.png
It is surprising that ANE ancestry today exists in Europe and the near east because there was little to no ANE ancestry in Loschbour and Stuttgart, there was some though in Motola12 estimated to be 19%.
I think migrations in the metal ages mainly or solely by Indo Europeans are why most modern Europeans have more WHG and ANE ancestry than Stuttgart and Loschbour did. That is especially evident in north-west Europeans aka Celto-Germanic people. WHG is 35% or more in Germanic(Norse-) people and Insular Celts, it is about 31% in French(not sure which region they are from but either way descended of Gauls), and is over 40% in Norse. Ancient genomes have proven north-west European farmers just 5,300 years ago where like Stuttgart and modern southwest Europeans.
WHG and ANE rose in north-west Europe starting around 5,000 years ago that is the same time Indo Europeans, and Y DNA R1b L11(dominate in west Europe) and R1a Z283(dominate in east Europe and popular in Scandinavia) arrived from eastern Europe. ANE is much lower in southwest Europe and could be almost completely absent in some. Alot of southwest Europe just 2,500 years ago was non Indo European the Romans brought Indo European language(Latin) to much of Spain, western France, and also brought it to Sardinia. Southwest Europeans happen to also be the closest relatives to Stuttgart and other Neolithic European samples I think this is more evidence WHG and ANE ancestry rose in north-west Europe because of Indo Europeans. WHG and ANE(in Europe) though may be highest in Uralic's who also have some ENA. This could be because Uralic's like Indo Europeans spread from far eastern Europe and because of farmer-hunter gatherer mixing.
French(aka Gauls) seem to be intermediate between Iberians(mixed Celtic, Aquitaine, and Iberian) and Germanics and Insular Celts. This could be because Indo European migrations came from the east and as they moved west they mixed more and more with the native Neolithic descended people. Maybe in central Europe(main R1b L11 branch is R1b S28) the Gauls had more WHG and ANE than Gauls in France especially southern and western France. It seems the Italic migration(almost all R1b L11 was R1b S28) to Italy from central Europe around 1,000BC did not raise WHG much but possibly rose ANE in Italy. I think extra near eastern ancestry was also brought to Italy and the Balkans after the Neolithic. More WHG in modern Europeans can also be from mixture between farmers and hunter gatherers that occurred during the Neolithic. There is evidence hunter gatherers were still in central Europe over 2,000 years after farmers had arrived(click here (http://www.sciencemag.org/content/342/6157/479.abstract)). I really doubt that explains all of the WHG most modern Europeans have that Neolithic farmer Stuttgart did not have. Laz 2013 thinks southern Europeans got their WHG mostly via EEF but northern Europeans got up to 50% additional WHG ancestry from another source I guess possibly farmer-hunter gatherer mixing that occurred after farming had established themselves.
I have heard some at Eurogenes hypothesis that early Indo Europeans(in far eastern Europe) had mainly ANE ancestry but also some WHG and EEF ancestry. Their reasoning is the even distribution of ANE in Europe, ANE in Europe is lowest in recently very non Indo European southwest Europe, and WHG's absence and ANE's existence in the near east even though there are Indo European languages in the near east. There is ANE in non Indo European near easterns so I don't think Indo Europeans can be the only source of ANE in the near east. The reason WHG seems to be absent in the near east could be because there is a small amount of ancestry from the people who brought Indo European languages to the near east.
mtDNA from bronze and iron age Indo Iranians show they had a very high amount of typical Mesolithic European mtDNA haplogroups U5, U4, and U2e. They also had majority light hair and eyes which today correlate very well with WHG so another sign of having a high amount of Mesolithic European ancestry. Indo European=ANE peoples counter argument to that is that U5, U4, and U2e also existed in ANE people(Loschbour had U5b1a and no ANE ancestry) which makes no sense to me and their other counter argument is that light hair and eyes existed in ANE people(Loschbour(no ANE ancestry) and La Brana-1 had the "blue eye" mutation) which also makes no sense to me how could ANE people develop the exact same mutations.
I think ANE in the near east probably has multiple sources the same could be true for ANE in Europe. It does seem based on ANE's distribution that Indo Europeans would have had a lot of it maybe more than WHG or EEF. I am assuming Indo Europeans were everything back then but really there were many different people that made migrations and extra ANE and WHG in modern Europeans may have a more complicated origin than Indo Europeans. ANE probably was brought to the near east by mainly or completely ANE people and some think early Indo Europeans were mainly ANE or completely ANE. I think know one really completely knows who the ANE people(s) were and how they contributed ancestry to so many modern people. Another reason why they may be connected to Indo Europeans is that Y DNA R1a1a1 M417 and R1b1a2a1a L11 are connected with Indo European migrations, ANE is base don MA1 who had Y DNA R(R1-, R2-), and native Americans who have the highest amount of ANE are entirely under Y DNA Q(specifically M3) which is the brotherclade to R.
I already made a thread about what I think of the pigmentation results from Laz 2013(Did Mesolithic Europeans had dark skin, dark hair, and blue eyes? (http://www.theapricity.com/forum/showthread.php?109997-Did-Mesolithic-Europeans-have-dark-skin-dark-hair-and-blue-eyes)). The "three light skin mutations" I don't think are the only factors to creating European light skin. The reason is all of them are about as popular in dark skinned near easterns and absent in Arabians who have basically the same skin color. The "three light skin mutations" I think may have been brought to Europe from the near east during the Neolithic or already were dominate Europe. There is no way Mesolithic Europeans had a high amount of blue eyes but dark skin and honestly I get very annoyed by people who stubbornly believe that's possible. WHG correlates very well with light hair, light skin, and light eyes in Europe today. I guess there could have been some Mesolithic Europeans who were dark(not a lot of blue eyes) and some that were light and it happens that the light ones blood survived much better after the Mesolithic. I doubt that though I think Mesolithic Europeans were light skinned and had multiple hair and eye colors. Modern day Sardinia people who are almost full blooded descendants of early Neolithic farmers like Stuttgart seems to have a mixture of light and dark pigmentation. I am basing this on Google images some can be as dark as near easterns and as light as northern Europeans. I think this is because of their mixed European hunter gatherer and near eastern farmer ancestry.
Finally here are the Y DNA, mtDNA, and pigmentation results from Laz 2013.
Loschbour, Heffingen Luxembourg (https://www.google.com/maps/preview/place/Heffingen,+Luxembourg/@49.768075,6.24983,13z/data=!3m1!4b1!4m2!3m1!1s0x47955989bb3c04bf:0x4f15b cdf4508e8c8) 6220-5990 BC mtDNA=U5b1a, Y DNA=I2a1b M423+ and L78+, I2a1b1 M359.2-, I2a1b2 L61.1-, I2a1b3 L621-. Ken Nordovt who was allowed to analyze SNP's from Loschbour thought to be phyloequivalent to M423 found that about half were derived and about half ancestral. So either pre-I2a1b(ancestral form of modern I2a1b) or from a related line to I2a1b that now is probably close to extinction.
Hair color:73.4% chance black hair, 25.6% chance brown hair, 1% chance blonde hair(97.5% chance of dark hair and 2.5% chance of light hair)
eye color:52.7% chance of blue eyes, 26.8% chance of intermediate, and 20.7% chance of brown eyes
skin color: had anything from light to dark brown. On the bases of the three so called light skin mutations he probably had some form of brown skin. There are probably other factors to European light skin that have not been discovered yet and he may have had them.
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f)6,000BC (https://www.google.com/maps/preview/place/Heffingen,+Luxembourg/@49.768075,6.24983,13z/data=!3m1!4b1!4m2!3m1!1s0x47955989bb3c04bf:0x4f15b cdf4508e8c8)Motola(1) mtDNA=U5a1
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(2) mtDNA=U2e1, Y DNA=I P38+ U179+ L41+, I1 M253-, I2a1b3 L621-, I2a2 L37-
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f)6,000BC Motola(3) mtDNA=U5a1, Y DNA=I2 L68+, I2a2 L181-, I2b L417-
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(4) mtDNA=U5a2d
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(6) mtDNA=U5a2d, Y DNA=Q1a2a L55+, but Q1 L232- so his Y DNA haplogroup is a mystery.
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(9) mtDNA=U5a2, Y DNA=I P38+, I1 P40-
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(12) mtDNA=U2e1, Y DNA=I2a1b* L78+, I2a1b1 M359.2-, I2a1b3 L621-. Ken Nordovt who was allowed to analyze SNP's from Motola12 thought to be phyloequivalent to M423 found that about half were derived and about half ancestral. So either pre-I2a1b(ancestral form of modern I2a1b) or from a related line to I2a1b that now is probably close to extinction.
Stuttgart, Germany LBK culture 7,500BP mtDNA=T2c1d1
hair color: 91.7% chance black, 7.9% chance brown, 0.4% chance blonde(99.8% chance dark and 0.2% chance light)
eye color: 99.4% chance brown and 0.6% chance intermediate
skin color: probably had light-olive skin
Y DNA I in all five of the Mesolithic samples is no surprise because Y DNA I has been theorized to be descended of pre Neolithic Europeans and today is very restricted to Europe like WHG ancestry. If tested I think all of the Y DNA samples from Motola, Sweden(except Motola6 who may have had Q1a2a L55) would have the same results Loschbour had so pre-I2a1b or a close relative to modern I2a1b. This probably means that lineage was widespread in Europe during the Mesolithic. Today I2a1b3 L621 is very popular in southeast Europe and I2a1b2 L161.1 is rare and is exclusively north-west European, and I don't know how I2a1b1 M359.2 is distributed. I2a1b may have formed in north-west Europe or relatives to Loschbour and Motola12's Y DNA somewhere else in Europe became modern I2a1b. This also probably means modern I2a1b subclades spread after the Mesolithic and with farmers.
Y DNA I1 takes up about 30-50% of modern Swedish Y DNA but it was not found in any of the Mesolithic Swedish I samples which were likely all pre-I2a1b or a close relative to modern I2a1b. Today I2a1 is around 1-5% in Sweden maybe less and on FTDNA I saw most of their I2a1 is M26- so I think most is probably I2a1b3 L621 or descended from the lineage the Mesolithic Motolas and Loschbour men were apart of.
It doesn't seem based on these samples that Mesolithic paternal lineages of west-central-north Europe have survived well. The likely true rumor that La Brana-1 a ~7,000 year old hunter gatherer from northern Spain had Y DNA C-V20 (http://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&cad=rja&sqi=2&ved=0CCcQFjAA&url=http%3A%2F%2Fwww.theapricity.com%2Fforum%2Fsho wthread.php%3F110804-Rearcher-posted-at-Eupedia-La-Brana-1-had-Y-DNA-C-V20&ei=8WzlUpfSA8iIqQGq64C4DQ&usg=AFQjCNFplJJBXlu_zDU78p0iPV43K9wVXg&sig2=IagtjRbC9lB4RX-a07OKGQ&bvm=bv.59930103,d.aWM) is even more evidence of this. I2a2, I1, I2a1(I2a1b3 L621-), and I2(I2a-) though likely descend from Mesolithic west-central-north Europeans. Today I2a1(I2a1b3 L621-) has a very exclusively western European distribution and probably descends from Mesolithic west Europeans. I2(I2a-) throughout Europe may have Mesolithic origin in what ever regions it is in. Y DNA I itself is estimated may be over 25,000 years old and probably originated in Europe but because there are many basal forms of I and I2 in the near east it may have originated there and quickly migrated to Europe in the upper Palaeolithic. C-V20 and F-96 are also likely haplogroups that have upper Palaeolithic European origins or at least existed in upper Palaeolithic Europe.
mtDNA U5 and U2e have already been found in Mesolithic Europe. Pre-U5 and U2(U2e- apart of another lineage) have been found in over 30,000 year old European mtDNA samples. Today U5 is very exclusive to Europe(like Y DNA I and WHG ancestry) and most diverse in Europe it probably arrived around 40,000 years ago from the near east as plain U* and developed into modern U5 by around 30,000 years ago. Most U2 subclades are mainly in south Asia except for U2e which I know takes up almost all U2e in Europe but I don't know about the near east. U4 was the second most popular mtDNA haplogroup of Mesolithic Europeans. Today it is most popular in Europe but not exclusive to Europe it also exists in the near east. I think mtDNA U5, U4, and U2e and Y DNA I, C-V20, and F-96 all dominated Mesolithic and upper Palaeolithic Europe, and that they all can be associated with WHG ancestry. That doesn't mean they all originated in Europe though.
All of the Swedish U5 samples were under U5a constant with all of the Neolithic Gotland hunter gatherers U5 samples with subclade being U5a. All pre historic U5 samples with subclade from Russia are also under U5a and under that U5a1. Russian hunter gatherers also had a high amount of U4 like the Gotland hunter gatherers and U2e like Motola hunter gatherers. So I think that may mean there was some type of genetic connection between Swedish and Russian Mesolithic hunter gatherers.
Analysis of ancient DNA can reveal historical events that are difficult to discern through study of present-day individuals. To investigate European population history around the time of the agricultural transition, we sequenced complete genomes from a ~7,500 year old early farmer from the Linearbandkeramik (LBK) culture from Stuttgart in Germany and an ~8,000 year old hunter-gatherer from the Loschbour rock shelter in Luxembourg. We also generated data from seven ~8,000 year old hunter-gatherers from Motala in Sweden. We compared these genomes and published ancient DNA to new data from 2,196 samples from 185 diverse populations to show that at least three ancestral groups contributed to present-day Europeans. The first are Ancient North Eurasians (ANE), who are more closely related to Upper Paleolithic Siberians than to any present-day population. The second are West European Hunter-Gatherers (WHG), related to the Loschbour individual, who contributed to all Europeans but not to Near Easterners. The third are Early European Farmers (EEF), related to the Stuttgart individual, who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model the deep relationships of these populations and show that about ~44% of the ancestry of EEF derived from a basal Eurasian lineage that split prior to the separation of other non-Africans.
If you want to download Lazaridis 2013 click here (http://biorxiv.org/content/early/2013/12/23/001552).
The first thread I made about Laz 2013(Y DNA I found in Mesolithic Sweden(I1-) and Luxemburg, a lot of other good stuff (http://www.theapricity.com/forum/showthread.php?107551-Y-DNA-I-found-in-Mesolithic-Sweden(-I1)-and-Luxemburg-alot-of-other-good-stuff)) was terrible because I posted it less than a day after Laz 2013 was published and I couldn't actually download the study so I got all of my information from posters. Even though now I have Laz 2013 downloaded I could study it for days, weeks, months, even years and still have many questions and unconfident conclusions. The most confusing thing to me are the ghost populations ANE and basal Eurasian. There are so many concepts involved with Laz 2013 that I can't remember it all in my head I have to write it down. The paper on La Brana-1's genome is suppose to be out tomorrow January 27th, 2014. I am sure the researchers compared La Brana-1's genome to data in Laz 2013, so it will help learn even more about European and overall human genetic history. It has been said by researcher Carles Lalueza-Fox La Brana-1 had the blue eye mutation(I assume it is the mutation in SNP rs12913832 (http://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=4&cad=rja&ved=0CEgQFjAD&url=http%3A%2F%2Fsnpedia.com%2Findex.php%2FRs12913 832&ei=QtjkUrWbJ8bhsAS4toKgCg&usg=AFQjCNHtVzdx9_RYDSyYm4KR8CKSXGWxkg&sig2=-fZWTGABJhLx4HQ9a-a4Ng&bvm=bv.59930103,d.aWM)) and Eupedia member CLF who claims to be Carles Lalueza-Fox said La Brana-1 had Y DNA C-V20 and that the paper should be out tomorrow.
Below are the three ancestral populations Laz 2013 claims modern Europeans descend from and the ancient individuals their based on, I coped and pasted this list from this (http://eurogenes.blogspot.com/2014/01/another-look-at-lazaridis-et-al-ancient.html) Eurogenes thread. The list includes ENA because north-east Europeans were shown to have some ENA ancestry(had already been discovered) also Sicilians, Maltese, and Ashkenazi Jews were found to maybe have some near eastern ancestry that can't only be explained by EEF.
- West European Hunter-Gatherer (WHG), based on an 8,000 year-old genome from Loschbour, Luxembourg
- Ancient North Eurasian (ANE), based on a 24,000 year-old genome from South Siberia (dubbed Mal'ta boy or MA-1)
- Early European Farmer (EEF), based on a 7,500 year-old genome from Stuttgart, Germany, belonging to the Neolithic Linearbandkeramik (LBK) culture
- Eastern non-African (ENA), this basically means East Eurasian, and is based on samples of present-day Onge, Han Chinese and Atayal from Taiwan
I have been confused who MA1 was genetically since the study that originally sampled his genome Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) was published in November 2013. In admixtures he looks like a mixture of European(specifically components that are most popular in north-eastern Europe and probably mainly of Mesolithic European origin), south-central Asian, native American, and maybe a little Oceania. I have been told by Davidski at Euorgenes that he is not a mix of those populations it is just that those components have MA1 like admixture.
Here are some important things I learned from Laz 2013 about the relationship between the ancient samples MA1, Loschbour, Motola12, and Stuttgart with eastern non Africans.
>Eurasian hunter gatherers(MA1, Stuttgart, and Motola12) are equally related to eastern non Africans,
>Eastern non Africans are more related to the Eurasian hunter gatherers(MA1, Loschbour, and Motola12) than to Stuttgart
>Eurasian hunter gatherers(MA1, Loschbour, and Motola12) and Stuttgart are more related to native Americans than they are to other eastern non Africans
>Native Americans are more related to MA1 than to European hunter gatherers(Loschbour and Motola12) and Stuttgart
>Papuans are less related to the Eurasian hunter gatherers(MA1, Loschbour, and Motola12) and Stuttgart than other eastern non Africans are Laz 2013 suggests that Denisovans admixture is the reason for this
>Atayal might be slightly less related to the ancient Eurasians(MA1, Loschbour, Motola12, and Stuttgart) than other eastern non Africans are.
Here is a figure from Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) "showing the levels of shared genetic drift between MA-1 and 147 present-day non-African populations", quote from Eurogenes thread First genome of an Upper Palaeolithic human. (http://eurogenes.blogspot.com/2013_11_01_archive.html)
http://img811.imageshack.us/img811/4151/zubo.png
MA1 is most related to native Americans(don't exactly know what shared drift means) but had no ENA ancestry. This means native Americans have some non ENA ancestry that is related to MA1. MA1 had mtDNA U(apart of his own distinct subclade) which is a west Eurasian mtDNA haplogroup(assumed, because it is so restricted to west Eurasians and most diverse in west Eurasians) he also had Y DNA R(R1-, R2-) which has a very debated origin, I don't think it originated in west Eurasians R more likely originated in a eastern non African people in my opinion because it descends from K-M526 and K-M256's except R and maybe Q(so their common ancestor P) originated in eastern non Africans. MA1 is 24,000 years old and R its self is estimated to be around 30,000 years old so maybe it originated in a people related to MA1 so maybe very related to west Eurasians and not eastern non African at all.
I don't know if MA1 was a west Eurasian person, from an extinct branch of the Eurasian family, or from a brother branch to west Eurasians. Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) says in their abstract there is evidence in MA1's autosomal DNA that he was basal to west Eurasians, closely related to native Americans, and had no affinity to east Asians. In PCA's MA1 and a 17,000 year old Siberian named AG2 clustered somewhat close to west Eurasians. Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) also said in their abstract that AG2 had similar autosomal genetic signatures as MA1. So I think MA1 was probably apart of a large population that was most related to west Eurasians and lived in Siberia(maybe other places) at least from 24,000-17,00 years ago and has left ancestry in many modern Eurasians and also in native Americans. I have noticed not many people consider that MA1 was a mutt. I think he may have had ancestry from many different(but related) populations that contributed ancestry to modern people separately.
Laz 2013 and Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) show that the relation to MA1 varies in west Eurasians which I think is prove there is also MA1 related ancestry in some west Eurasians. Relatedness to MA1 in the ancient European samples also varied because Stuttgart and Loschbour had little to no MA1 related ancestry but MA1 shared more alleles with Motola12 than he did with Loschbour. It was estimated that Motola12 had 81% Loschbour related ancestry and 19% MA1 related ancestry.
Laz 2013 found in many different tests that all modern Europeans(except maybe Basque, French_South, Bergamo, Pais_Vasco, Sardinian) are not an admixture of just Loschbour and Stuttgart related populations but also have MA1 related ancestry. Some Europeans scored a lower F3-statistic with Stuttgart-MA1 than Stuttgart-Loschbour showing some Europeans have more relation to MA1 than Loschbour and evidence for MA1 related ancestry. The F4-statistic(test population, Stuttgart, MA1, Chimp) is positive in many Europeans and near easterns but when MA1 is replaced by Loschbour it is only positive in Europeans. Meaning Loschbour related ancestry may be restricted to Europe but MA1 and Stuttgart related ancestry exists in Europe and the near east.
In F-3 statistics that included every possible modern west Eurasian population with at least four samples and one of the ancient samples as a at least one of the references, every near eastern population had its lowest statistic with Stuttgart as one of the references none had Loschbour or Motola12 as one of the references. In a F4 statistic it was shown that most modern Europeans have more Loschbour related ancestry than Stuttgart does but it also showed all modern near easterns have less Loschbour related ancestry than Stuttgart had. Evidence there is no close relationship between Loschbour and modern near easterns, Stuttgart derived from mainly the same source that modern near easterns derive from, and that Stuttgart had Loschbour related ancestry it was predicted that she had less than 100% near eastern ancestry and possibly as little as 61%.
I am very surprised by how unrelated modern near easterns are to Loschbour because in many admixtures like globe13 the component(s) that are most popular in north-eastern Europe and are mainly Mesolithic Europeans descended are the closest relative(s) to the component(s) that are most popular in northern west Asia. Since the similar results came from so many different admixtures I thought there must be some type of close relationship between Mesolithic Europeans and northern west Asians.
I am also very surprised by how related Stuttgart is to modern near easterns and that near easterns all probably descend from the same source. This is exactly what many people assumed though because farming spread to Europe from the near east. mtDNA and Y DNA of Neolithic European farmers does show that they were more related paternally and maternally to modern near easterns than any modern near easterns are to Mesolithic Europeans. It would make more sense to me that the near east is very diverse with many different west Eurasian family groups but Laz 2013 gives evidence they all come from the same root and that European(Loschbour) is their brother clade.
I really don't understand why Stuttgart would have ~44% basal Eurasian ancestry, from a people that separated from the Eurasian root before the east and west split. If this is true that means there is a large amount of basal Eurasian ancestry in Europeans, near easterns, and probably north Africans. I guess basal Eurasian ancestry could be an explanation of why Stuttgart is less related to eastern non Africans than ancient Eurasian hunter gatherers(MA1, Loschbour, and Motola12) were. I haven't seen anyone question how basal Eurasian ancestry would affect looks. I doubt Sardinians and Stuttgart would have looked like typical west Eurasians(which they do) with having ~44% non west Eurasian ancestry. If the basal Eurasian thing is somewhat true then I think possibly Y DNA E1b1b and G2a and mtDNA N1a1a and W in Neolithic Europeans may be connected.
No First I will say for native Americans there is no EEF or WHG ancestry just ANE which was predicted by Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) to be about 14-38% of their total ancestry.
Laz 2013 made an admixture test EEF(Stuttgart), WHG(Loschbour), and ANE(MA1) it really only works for Europeans though and for some Europeans who have extra near eastern ancestry or east Asian ancestry the results will be inaccurate. Eurogenes project allows members who have Eurogenes K13 results(click here (http://bga101.blogspot.com/2013/12/eef-whg-ane-test-for-europeans.html)) to get results in this test.
Iberia and France
<tbody>
Population
EEF
WHG
ANE
Basque
59.3
29.3
11.4
Pais_Vasco
71.3
12.5
16.3
Spanish
80.9
6.8
12.3
French
55.4
31.1
13.5
French_South
67.5
19.5
13
</tbody>
North west Europe
<tbody>
Population
EEF
WHG
ANE
Icelandic
39.4
45.6
15
English
49.5
36.4
14.1
Norwegian
41.1
42.8
16.1
Orcadian
45.7
38.5
15.8
Scottish
39
42.8
18.2
</tbody>
North east Europe
<tbody>
Population
EEF
WHG
ANE
Lithuanian
36.4
46.4
17.2
Estonian
32.2
49.5
18.3
</tbody>
Slavic east Europe(X Balkans)
<tbody>
Population
EEF
WHG
ANE
Ukrainian
46.2
38.7
15.1
Belorussian
41.8
43.1
15.1
Czech
49.5
33.8
16.7
</tbody>
Balkans an Italy
<tbody>
Population
EEF
WHG
ANE
Bergamo
71.5
17.7
10.8
Bulgarian
71.2
14.7
14.1
Croatian
56.1
29.3
14.5
Greek
79.2
5.8
15.1
Tuscan
74.6
13.6
11.8
Sardinian
81.7
17.5
0.8
Sicilian
90.3
0
9.7
Albanian
78.1
9.2
12.7
</tbody>
Near east
<tbody>
Near east
EEF
WHG
ANE
Ashkenazi_Jew
93.1
0
6.9
Maltese
93.2
0
6.8
</tbody>
Laz 2013 like many other people have found that Loschbour(or European hunter gatherers in general) are most related to modern northern Europeans especially north eastern and Scandinavian and that Stuttgart(or Neolithic European farmers in general) are more related to modern southern Europeans especially southwestern like Basque and Sardinia. In the PCA's Sardinians cluster very closely to Stuttgart and in K=2-K=20 admixtures from Laz 2013 had nearly identical results like they did with Otzi(~5,300BP early copper age farmer from the alps) and ~5,000 year old Swedish Funnel beaker farmers. I think Sardinians are nearly completely un mixed descendants of Stuttgart like farmers and during the Neolithic all central-west-northern(maybe other regions) European farmers were Stuttgart/Sardinia like.
The distribution of WHG ancestry is nearly identical to component(s) in many different admixtures usually called north European or another name referring to an area in northern Europe. The modern componets though probably have some non European hunter gatherer admixture. When near easterns are put in this admixture EEF, WHG, and ANE they usually have over 100% EEF and negative WHG this is constant with other evidence that seem to say there is little to no WHG ancestry in the near east.
ANE unlike WHG and EEF has a pretty even distribution in Europe. It seems to be a little bit higher in northern Europe though especially in north-east Europeans. ANE ancestry though is probably by far highest in native Americans and is also probably higher in the Caucasus than anywhere in Europe. When put in an admixture with two options near east and ANE Caucasus populations Abkhasian scored 18.6% ANE, Chechen scored 27%, and Lezgin scored 28.8%. There is also scattered ANE ancestry in the near east but at a lower levels than in what most Europeans have.
This is a map of the figure from Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013) from the begging of this thread which shows that MA1 is more related to Europeans than near easterns, and there is a PCA from Raghavan 2013 (http://www.ancestraljourneys.org/bibliography.shtml#Raghavan2013).
http://3.bp.blogspot.com/-qdDK2ZHSeV4/UpxqAm3iUvI/AAAAAAAACRY/pthyx4u0VzU/s1600/Mal%27ta1.png
It is surprising that ANE ancestry today exists in Europe and the near east because there was little to no ANE ancestry in Loschbour and Stuttgart, there was some though in Motola12 estimated to be 19%.
I think migrations in the metal ages mainly or solely by Indo Europeans are why most modern Europeans have more WHG and ANE ancestry than Stuttgart and Loschbour did. That is especially evident in north-west Europeans aka Celto-Germanic people. WHG is 35% or more in Germanic(Norse-) people and Insular Celts, it is about 31% in French(not sure which region they are from but either way descended of Gauls), and is over 40% in Norse. Ancient genomes have proven north-west European farmers just 5,300 years ago where like Stuttgart and modern southwest Europeans.
WHG and ANE rose in north-west Europe starting around 5,000 years ago that is the same time Indo Europeans, and Y DNA R1b L11(dominate in west Europe) and R1a Z283(dominate in east Europe and popular in Scandinavia) arrived from eastern Europe. ANE is much lower in southwest Europe and could be almost completely absent in some. Alot of southwest Europe just 2,500 years ago was non Indo European the Romans brought Indo European language(Latin) to much of Spain, western France, and also brought it to Sardinia. Southwest Europeans happen to also be the closest relatives to Stuttgart and other Neolithic European samples I think this is more evidence WHG and ANE ancestry rose in north-west Europe because of Indo Europeans. WHG and ANE(in Europe) though may be highest in Uralic's who also have some ENA. This could be because Uralic's like Indo Europeans spread from far eastern Europe and because of farmer-hunter gatherer mixing.
French(aka Gauls) seem to be intermediate between Iberians(mixed Celtic, Aquitaine, and Iberian) and Germanics and Insular Celts. This could be because Indo European migrations came from the east and as they moved west they mixed more and more with the native Neolithic descended people. Maybe in central Europe(main R1b L11 branch is R1b S28) the Gauls had more WHG and ANE than Gauls in France especially southern and western France. It seems the Italic migration(almost all R1b L11 was R1b S28) to Italy from central Europe around 1,000BC did not raise WHG much but possibly rose ANE in Italy. I think extra near eastern ancestry was also brought to Italy and the Balkans after the Neolithic. More WHG in modern Europeans can also be from mixture between farmers and hunter gatherers that occurred during the Neolithic. There is evidence hunter gatherers were still in central Europe over 2,000 years after farmers had arrived(click here (http://www.sciencemag.org/content/342/6157/479.abstract)). I really doubt that explains all of the WHG most modern Europeans have that Neolithic farmer Stuttgart did not have. Laz 2013 thinks southern Europeans got their WHG mostly via EEF but northern Europeans got up to 50% additional WHG ancestry from another source I guess possibly farmer-hunter gatherer mixing that occurred after farming had established themselves.
I have heard some at Eurogenes hypothesis that early Indo Europeans(in far eastern Europe) had mainly ANE ancestry but also some WHG and EEF ancestry. Their reasoning is the even distribution of ANE in Europe, ANE in Europe is lowest in recently very non Indo European southwest Europe, and WHG's absence and ANE's existence in the near east even though there are Indo European languages in the near east. There is ANE in non Indo European near easterns so I don't think Indo Europeans can be the only source of ANE in the near east. The reason WHG seems to be absent in the near east could be because there is a small amount of ancestry from the people who brought Indo European languages to the near east.
mtDNA from bronze and iron age Indo Iranians show they had a very high amount of typical Mesolithic European mtDNA haplogroups U5, U4, and U2e. They also had majority light hair and eyes which today correlate very well with WHG so another sign of having a high amount of Mesolithic European ancestry. Indo European=ANE peoples counter argument to that is that U5, U4, and U2e also existed in ANE people(Loschbour had U5b1a and no ANE ancestry) which makes no sense to me and their other counter argument is that light hair and eyes existed in ANE people(Loschbour(no ANE ancestry) and La Brana-1 had the "blue eye" mutation) which also makes no sense to me how could ANE people develop the exact same mutations.
I think ANE in the near east probably has multiple sources the same could be true for ANE in Europe. It does seem based on ANE's distribution that Indo Europeans would have had a lot of it maybe more than WHG or EEF. I am assuming Indo Europeans were everything back then but really there were many different people that made migrations and extra ANE and WHG in modern Europeans may have a more complicated origin than Indo Europeans. ANE probably was brought to the near east by mainly or completely ANE people and some think early Indo Europeans were mainly ANE or completely ANE. I think know one really completely knows who the ANE people(s) were and how they contributed ancestry to so many modern people. Another reason why they may be connected to Indo Europeans is that Y DNA R1a1a1 M417 and R1b1a2a1a L11 are connected with Indo European migrations, ANE is base don MA1 who had Y DNA R(R1-, R2-), and native Americans who have the highest amount of ANE are entirely under Y DNA Q(specifically M3) which is the brotherclade to R.
I already made a thread about what I think of the pigmentation results from Laz 2013(Did Mesolithic Europeans had dark skin, dark hair, and blue eyes? (http://www.theapricity.com/forum/showthread.php?109997-Did-Mesolithic-Europeans-have-dark-skin-dark-hair-and-blue-eyes)). The "three light skin mutations" I don't think are the only factors to creating European light skin. The reason is all of them are about as popular in dark skinned near easterns and absent in Arabians who have basically the same skin color. The "three light skin mutations" I think may have been brought to Europe from the near east during the Neolithic or already were dominate Europe. There is no way Mesolithic Europeans had a high amount of blue eyes but dark skin and honestly I get very annoyed by people who stubbornly believe that's possible. WHG correlates very well with light hair, light skin, and light eyes in Europe today. I guess there could have been some Mesolithic Europeans who were dark(not a lot of blue eyes) and some that were light and it happens that the light ones blood survived much better after the Mesolithic. I doubt that though I think Mesolithic Europeans were light skinned and had multiple hair and eye colors. Modern day Sardinia people who are almost full blooded descendants of early Neolithic farmers like Stuttgart seems to have a mixture of light and dark pigmentation. I am basing this on Google images some can be as dark as near easterns and as light as northern Europeans. I think this is because of their mixed European hunter gatherer and near eastern farmer ancestry.
Finally here are the Y DNA, mtDNA, and pigmentation results from Laz 2013.
Loschbour, Heffingen Luxembourg (https://www.google.com/maps/preview/place/Heffingen,+Luxembourg/@49.768075,6.24983,13z/data=!3m1!4b1!4m2!3m1!1s0x47955989bb3c04bf:0x4f15b cdf4508e8c8) 6220-5990 BC mtDNA=U5b1a, Y DNA=I2a1b M423+ and L78+, I2a1b1 M359.2-, I2a1b2 L61.1-, I2a1b3 L621-. Ken Nordovt who was allowed to analyze SNP's from Loschbour thought to be phyloequivalent to M423 found that about half were derived and about half ancestral. So either pre-I2a1b(ancestral form of modern I2a1b) or from a related line to I2a1b that now is probably close to extinction.
Hair color:73.4% chance black hair, 25.6% chance brown hair, 1% chance blonde hair(97.5% chance of dark hair and 2.5% chance of light hair)
eye color:52.7% chance of blue eyes, 26.8% chance of intermediate, and 20.7% chance of brown eyes
skin color: had anything from light to dark brown. On the bases of the three so called light skin mutations he probably had some form of brown skin. There are probably other factors to European light skin that have not been discovered yet and he may have had them.
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f)6,000BC (https://www.google.com/maps/preview/place/Heffingen,+Luxembourg/@49.768075,6.24983,13z/data=!3m1!4b1!4m2!3m1!1s0x47955989bb3c04bf:0x4f15b cdf4508e8c8)Motola(1) mtDNA=U5a1
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(2) mtDNA=U2e1, Y DNA=I P38+ U179+ L41+, I1 M253-, I2a1b3 L621-, I2a2 L37-
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f)6,000BC Motola(3) mtDNA=U5a1, Y DNA=I2 L68+, I2a2 L181-, I2b L417-
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(4) mtDNA=U5a2d
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(6) mtDNA=U5a2d, Y DNA=Q1a2a L55+, but Q1 L232- so his Y DNA haplogroup is a mystery.
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(9) mtDNA=U5a2, Y DNA=I P38+, I1 P40-
Motola, Sweden (https://www.google.com/maps/preview/place/Motala,+Sweden/@58.5483891,15.0420942,12z/data=!3m1!4b1!4m2!3m1!1s0x465bdcd950a4d43d:0x1fd1c 7707f1ccb4f) 6,000BC Motola(12) mtDNA=U2e1, Y DNA=I2a1b* L78+, I2a1b1 M359.2-, I2a1b3 L621-. Ken Nordovt who was allowed to analyze SNP's from Motola12 thought to be phyloequivalent to M423 found that about half were derived and about half ancestral. So either pre-I2a1b(ancestral form of modern I2a1b) or from a related line to I2a1b that now is probably close to extinction.
Stuttgart, Germany LBK culture 7,500BP mtDNA=T2c1d1
hair color: 91.7% chance black, 7.9% chance brown, 0.4% chance blonde(99.8% chance dark and 0.2% chance light)
eye color: 99.4% chance brown and 0.6% chance intermediate
skin color: probably had light-olive skin
Y DNA I in all five of the Mesolithic samples is no surprise because Y DNA I has been theorized to be descended of pre Neolithic Europeans and today is very restricted to Europe like WHG ancestry. If tested I think all of the Y DNA samples from Motola, Sweden(except Motola6 who may have had Q1a2a L55) would have the same results Loschbour had so pre-I2a1b or a close relative to modern I2a1b. This probably means that lineage was widespread in Europe during the Mesolithic. Today I2a1b3 L621 is very popular in southeast Europe and I2a1b2 L161.1 is rare and is exclusively north-west European, and I don't know how I2a1b1 M359.2 is distributed. I2a1b may have formed in north-west Europe or relatives to Loschbour and Motola12's Y DNA somewhere else in Europe became modern I2a1b. This also probably means modern I2a1b subclades spread after the Mesolithic and with farmers.
Y DNA I1 takes up about 30-50% of modern Swedish Y DNA but it was not found in any of the Mesolithic Swedish I samples which were likely all pre-I2a1b or a close relative to modern I2a1b. Today I2a1 is around 1-5% in Sweden maybe less and on FTDNA I saw most of their I2a1 is M26- so I think most is probably I2a1b3 L621 or descended from the lineage the Mesolithic Motolas and Loschbour men were apart of.
It doesn't seem based on these samples that Mesolithic paternal lineages of west-central-north Europe have survived well. The likely true rumor that La Brana-1 a ~7,000 year old hunter gatherer from northern Spain had Y DNA C-V20 (http://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&cad=rja&sqi=2&ved=0CCcQFjAA&url=http%3A%2F%2Fwww.theapricity.com%2Fforum%2Fsho wthread.php%3F110804-Rearcher-posted-at-Eupedia-La-Brana-1-had-Y-DNA-C-V20&ei=8WzlUpfSA8iIqQGq64C4DQ&usg=AFQjCNFplJJBXlu_zDU78p0iPV43K9wVXg&sig2=IagtjRbC9lB4RX-a07OKGQ&bvm=bv.59930103,d.aWM) is even more evidence of this. I2a2, I1, I2a1(I2a1b3 L621-), and I2(I2a-) though likely descend from Mesolithic west-central-north Europeans. Today I2a1(I2a1b3 L621-) has a very exclusively western European distribution and probably descends from Mesolithic west Europeans. I2(I2a-) throughout Europe may have Mesolithic origin in what ever regions it is in. Y DNA I itself is estimated may be over 25,000 years old and probably originated in Europe but because there are many basal forms of I and I2 in the near east it may have originated there and quickly migrated to Europe in the upper Palaeolithic. C-V20 and F-96 are also likely haplogroups that have upper Palaeolithic European origins or at least existed in upper Palaeolithic Europe.
mtDNA U5 and U2e have already been found in Mesolithic Europe. Pre-U5 and U2(U2e- apart of another lineage) have been found in over 30,000 year old European mtDNA samples. Today U5 is very exclusive to Europe(like Y DNA I and WHG ancestry) and most diverse in Europe it probably arrived around 40,000 years ago from the near east as plain U* and developed into modern U5 by around 30,000 years ago. Most U2 subclades are mainly in south Asia except for U2e which I know takes up almost all U2e in Europe but I don't know about the near east. U4 was the second most popular mtDNA haplogroup of Mesolithic Europeans. Today it is most popular in Europe but not exclusive to Europe it also exists in the near east. I think mtDNA U5, U4, and U2e and Y DNA I, C-V20, and F-96 all dominated Mesolithic and upper Palaeolithic Europe, and that they all can be associated with WHG ancestry. That doesn't mean they all originated in Europe though.
All of the Swedish U5 samples were under U5a constant with all of the Neolithic Gotland hunter gatherers U5 samples with subclade being U5a. All pre historic U5 samples with subclade from Russia are also under U5a and under that U5a1. Russian hunter gatherers also had a high amount of U4 like the Gotland hunter gatherers and U2e like Motola hunter gatherers. So I think that may mean there was some type of genetic connection between Swedish and Russian Mesolithic hunter gatherers.