Admixture between early LBK and HGs

Angela

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See:
Alexei Nikitin et al (plus Reich and Lazaridis)
https://www.biorxiv.org/content/biorxiv/early/2019/08/21/741900.full.pdf
[h=1]"Interactions between earliest Linearbandkeramik farmers and central European hunter gatherers at the dawn of European Neolithization"[/h]
"[FONT=&quot]Archaeogenetic research over the last decade has demonstrated that European Neolithic farmers (ENFs) were descended primarily from Anatolian Neolithic farmers (ANFs). ENFs, including early Neolithic central European [/FONT]Linearbandkeramik[FONT=&quot] (LBK) farming communities, also harbored ancestry from European Mesolithic hunter gatherers (WHGs) to varying extents, reflecting admixture between ENFs and WHGs. However, the timing and other details of this process are still imperfectly understood. In this report, we provide a bioarchaeological analysis of three individuals interred at the Brunn 2 site of the Brunn am Gebirge-Wolfholz archeological complex, one of the oldest LBK sites in central Europe. Two of the individuals had a mixture of WHG-related and ANF-related ancestry, one of them with approximately 50% of each, while the third individual had approximately all ANF-related ancestry. Stable carbon and nitrogen isotope ratios for all three individuals were within the range of variation reflecting diets of other Neolithic agrarian populations. Strontium isotope analysis revealed that the ~50% WHG-ANF individual was non-local to the Brunn 2 area. Overall, our data indicate interbreeding between incoming farmers, whose ancestors ultimately came from western Anatolia, and local HGs, starting within the first few generations of the arrival of the former in central Europe, as well as highlighting the integrative nature and composition of the early LBK communities."

[/FONT]
I think they're probably right, but I'm not sure three samples are enough for conclusive findings.

It's interesting to see how spotty it was, with one sample having no HG, one being half and half, and one having some HG. Of course, the eventual outcome was a group with not very much HG, not like MN farmers.

It's a lesson in not being too dogmatic about long term effects on the genome of an area of some admixture results from an early stage and one area.

 
The presentation and discussion of articles like this are why I remain with eupedia.com. I look forward to the commentary to come (I'm truly not qualified to comment).
 
afaik LBK farmers had the least WHG admixture - some 5%

That's my understanding too, but we only know that because we have a lot of WHG samples, and about 5% is what it averages out to be.

If this group of LBK people were the first ones we found, we might have been led to believe that there was a lot of admixture in this very early Neolithic group, while it's actually not the case at all.

That has implications for other papers, and the perhaps hasty decisions we reach about the nature of people in certain areas and time periods. Think of the Parma North Italian Beakers. We have, what, three of them? How representative of the people of Northern Italy at that time will they turn out to be?

https://www.eupedia.com/forum/threa...orthwest-Europe/page4?highlight=Parma+Beakers
 
LBK farmers and European hunter gatherers at the dawn of European Neolithization

https://www.nature.com/articles/s41598-019-56029-2

Abstract


Archaeogenetic research over the last decade has demonstrated that European Neolithic farmers (ENFs) were descended primarily from Anatolian Neolithic farmers (ANFs). ENFs, including early Neolithic central European Linearbandkeramik (LBK) farming communities, also harbored ancestry from European Mesolithic hunter gatherers (WHGs) to varying extents, reflecting admixture between ENFs and WHGs. However, the timing and other details of this process are still imperfectly understood. In this report, we provide a bioarchaeological analysis of three individuals interred at the Brunn 2 site of the Brunn am Gebirge-Wolfholz archeological complex, one of the oldest LBK sites in central Europe. Two of the individuals had a mixture of WHG-related and ANF-related ancestry, one of them with approximately 50% of each, while the third individual had approximately all ANF-related ancestry. Stable carbon and nitrogen isotope ratios for all three individuals were within the range of variation reflecting diets of other Neolithic agrarian populations. Strontium isotope analysis revealed that the ~50% WHG-ANF individual was non-local to the Brunn 2 area. Overall, our data indicate interbreeding between incoming farmers, whose ancestors ultimately came from western Anatolia, and local HGs, starting within the first few generations of the arrival of the former in central Europe, as well as highlighting the integrative nature and composition of the early LBK communities.
 
https://www.nature.com/articles/s41598-019-56029-2

Abstract


Archaeogenetic research over the last decade has demonstrated that European Neolithic farmers (ENFs) were descended primarily from Anatolian Neolithic farmers (ANFs). ENFs, including early Neolithic central European Linearbandkeramik (LBK) farming communities, also harbored ancestry from European Mesolithic hunter gatherers (WHGs) to varying extents, reflecting admixture between ENFs and WHGs. However, the timing and other details of this process are still imperfectly understood. In this report, we provide a bioarchaeological analysis of three individuals interred at the Brunn 2 site of the Brunn am Gebirge-Wolfholz archeological complex, one of the oldest LBK sites in central Europe. Two of the individuals had a mixture of WHG-related and ANF-related ancestry, one of them with approximately 50% of each, while the third individual had approximately all ANF-related ancestry. Stable carbon and nitrogen isotope ratios for all three individuals were within the range of variation reflecting diets of other Neolithic agrarian populations. Strontium isotope analysis revealed that the ~50% WHG-ANF individual was non-local to the Brunn 2 area. Overall, our data indicate interbreeding between incoming farmers, whose ancestors ultimately came from western Anatolia, and local HGs, starting within the first few generations of the arrival of the former in central Europe, as well as highlighting the integrative nature and composition of the early LBK communities.

Thanks, Jose, but we already have a thread on it, with some discussion.

See:
https://www.eupedia.com/forum/threa...nd-HGs?highlight=dawn+European+Neolithization
 
DNA from a man who lived in Ethiopia about 4,500 years ago is prompting scientists to rethink the history of human migration in Africa.
Until now, the conventional wisdom had been that the first groups of modern humans left Africa roughly 70,000 years ago, stopping in the Middle East en route to Europe, Asia and beyond. Then about 3,000 years ago, a group of farmers from the Middle East and present-day Turkey came back to the Horn of Africa (probably bringing crops like wheat, barley and lentils with them).

Population geneticists pieced this story together by comparing the DNA of distinct groups of people alive today. Since humans emerged in Africa, DNA from an ancient Africa could provide a valuable genetic baseline that would make it easier for scientists to track genome changes over timesUnfortunately, such DNA has been hard to come by. DNA isn’t built to last for thousands of years. The samples of ancient DNA that have been sequenced to date were extracted from bodies in Europe and Asia that were naturally refrigerated in cooler climates.

That’s what makes the Ethiopian man so special. His body was found face-down in Mota cave, which is situated in the highlands in the southern part of the country. The cool, dry conditions in the cave preserved his DNA, and scientists extracted a sample from the petrous bone at the base of his skull. The resulting sequence is the first nuclear genome from an ancient African, according to a report published Thursday in the journal Science.

Radiocarbon dating revealed that the bone was 4,500 years old. That meant Mota (as the researchers called him) lived before Eurasians returned to the African continent.


Consistent with that timeline, Mota did not have any of the genetic variants for light-colored eyes or skin that evolved in the populations that left Africa. Nor did he have variants that arose in Eurasian farmers that allowed them to digest milk as adults.



Mota did have three variants that are known to help modern-day Ethiopians live in high altitudes. (The present-day town of Mota lies more than 8,100 feet above sea level.)

When the researchers compared Mota’s genome to those of contemporary humans, the closest match was with the Ari people of southern Ethiopia.

With this information, the research team was able to investigate the mysterious group of Eurasians that came to Africa 3,000 years ago. They created a model that assumed the Ari genome was a mixture of DNA from Mota and an unknown population from west Eurasia. Then they “plugged in” DNA from several candidate populations to see if they could get a combination that looked like Ari DNA.

Two results stood out from the rest. One was for modern-day Sardinians, who are known to be the closest living relatives to the earliest farmers. The other was for members of the so-called LBK culture in Germany, early farmers who lived about 7,000 years ago.


If the Eurasian settlers who arrived in Africa 3,000 years ago were indeed descendants of the LBK farmers, then the story of their migration through Africa needs to be revised, the researchers wrote.


By comparing the LBK genome with DNA from Africans alive today, the scientists calculated that these ancient farmers may have made up 25% or more of the population in the Horn of Africa during the migration years. All of those migrants ultimately pushed farther into Africa than previously thought, they determined.


African populations from the western and southern tips of the continent got at least 5% of their DNA from these Eurasian migrants, according to the study. Some groups from Ethiopia, Somalia, Djibouti and Eritrea can trace more than 30% of their DNA to these migrants.


“The ability to sequence ancient genomes has revolutionized our understanding of human evolution,” wrote the research team, which was led by Marcos Gallego Llorente of the University of Cambridge and Eppie Ruth Jones of Trinity College Dublin. They said they are eager to find “even older African genomes” that may make the story more complete.

https://www.latimes.com/science/sci...ent-ethiopian-dna-eurasia-20151008-story.html
 
Thank you so much Mrs. Angela. If it were possible I would like to eliminate this thread. As you are a prominent person at Eupedia you may be able to help to amend this: At Eupedia, not in this chapter of the forum, but in the genetics chapter, in the Distribution maps of autosomal admixtures in Europe, the Middle East and North Africa, in the text of the Early European Farmer (EEF) map admixture states: The closest modern populations of the EEF are the Ashkenazi Jews (93%). I have an idea this is between 40 and 50%. On the map itself, in the territory of Israel, the colors are in agreement with 93%. Sardinia is less loaded than Secilia, is that correct?
 
Thank you so much Mrs. Angela. If it were possible I would like to eliminate this thread. As you are a prominent person at Eupedia you may be able to help to amend this: At Eupedia, not in this chapter of the forum, but in the genetics chapter, in the Distribution maps of autosomal admixtures in Europe, the Middle East and North Africa, in the text of the Early European Farmer (EEF) map admixture states: The closest modern populations of the EEF are the Ashkenazi Jews (93%). I have an idea this is between 40 and 50%. On the map itself, in the territory of Israel, the colors are in agreement with 93%. Sardinia is less loaded than Secilia, is that correct?

I think some of that material is dated, Jose. Also, those particular maps used amateur data from Eurogenes. The papers show a different story. Then there slight differences from paper to paper, depending on which Early Neolithic sample they're using.

This is pretty reliable, however.

F3.large.jpg


The Neolithic is on the left, the steppe admixed Bronze Age populations on the right. If you're looking at the Cardial like EEF, Spain is second to Sardinia. If you're looking at Danubian EEF, Sicily and Northern Italy are perhaps even a bit higher than Sardinia. So, it depends.

You can look at it in this Haak et al chart too, which I like because it's not based on Admixture, but on more sophisticated statistics. This one is using a sample from LBK, so earlier, less WHG.
Haak-et-al-2015-Figure-3-Admixture-Proportions-in-Modern-DNA-With-Linguistic-and-Historical-Origins-Added.png



As the first graphic showed, nobody in the modern Near East has anywhere near the amount of EEF that is in Europeans. That's bkz they got so much Caucasus/Iranian like input after the farmers left for Europe.

Ashkenazi Jews are somewhere in the general vicinity of Sicily and Southern Italy, so that will give you an idea. The Judeans clearly picked up a lot of ancestry in Europe. Until we have an ancient sample from Judea in, say, the time of Maccabees, we won't know how much.
 
I think some of that material is dated, Jose. Also, those particular maps used amateur data from Eurogenes. The papers show a different story. Then there slight differences from paper to paper, depending on which Early Neolithic sample they're using.

This is pretty reliable, however.

F3.large.jpg


The Neolithic is on the left, the steppe admixed Bronze Age populations on the right. If you're looking at the Cardial like EEF, Spain is second to Sardinia. If you're looking at Danubian EEF, Sicily and Northern Italy are perhaps even a bit higher than Sardinia. So, it depends.

You can look at it in this Haak et al chart too, which I like because it's not based on Admixture, but on more sophisticated statistics. This one is using a sample from LBK, so earlier, less WHG.
Haak-et-al-2015-Figure-3-Admixture-Proportions-in-Modern-DNA-With-Linguistic-and-Historical-Origins-Added.png



As the first graphic showed, nobody in the modern Near East has anywhere near the amount of EEF that is in Europeans. That's bkz they got so much Caucasus/Iranian like input after the farmers left for Europe.

Ashkenazi Jews are somewhere in the general vicinity of Sicily and Southern Italy, so that will give you an idea. The Judeans clearly picked up a lot of ancestry in Europe. Until we have an ancient sample from Judea in, say, the time of Maccabees, we won't know how much.

How does Spain show zero WHG in the Haak chart? They have a significant amount of mtDNA U5 and y-dna I2.
 
How does Spain show zero WHG in the Haak chart? They have a significant amount of mtDNA U5 and y-dna I2.

You'd have to check me, but to the best of my recollection most of that is in the Spanish Basques, whose results you can see labelled as "Spanish North".
 
You'd have to check me, but to the best of my recollection most of that is in the Spanish Basques, whose results you can see labelled as "Spanish North".

Region/Haplogroup
[TH="class: I1"]I1[/TH]
[TH="class: I2"]I2*/I2a[/TH]
[TH="class: I2b"]I2b[/TH]
[TH="class: R1a"]R1a[/TH]
[TH="class: R1b"]R1b[/TH]
[TH="class: G2a"]G[/TH]
[TH="class: J2"]J2[/TH]
[TH="class: J1"]J*/J1[/TH]
[TH="class: E1b"]E1b1b[/TH]
[TH="class: T"]T[/TH]
[TH="class: Q"]Q[/TH]
[TH="class: N1c1"]N[/TH]
[TD="class: samples"]Sample size[/TD]
Spain1.54.51269 381.572.500
rate11.gif
Andalusia09.503.558.5310.5210300
rate07.gif
Aragon214.51260.5110.505400
rate05.gif
Asturias2202.558.588214300
rate05.gif
Basque country0.5500851.52.50.52.500.50
rate07.gif
Cantabria1328.55510.532.5112.500
rate05.gif
Castile & Leon0.520.536456116200
rate07.gif
Castile-La-Mancha1.51.50.51.56681045200
rate05.gif
Catalonia23.51.51.566.54.57.51.58.5100
rate11.gif
Extremadura3.551050511.5018.5500
rate05.gif
Galicia53.52157410.5410.5200
rate07.gif

Region/Haplogroup
[TD="class: mL"]L[/TD]
[TD="class: HV"]HV[/TD]
[TD="class: H"]H[/TD]
[TD="class: H, width: 40"]H1+H3[/TD]
[TD="class: H, width: 40"]H5[/TD]
[TD="class: V, width: 40"]HV0+V[/TD]
[TD="class: J"]J[/TD]
[TD="class: mT"]T1[/TD]
[TD="class: mT"]T2[/TD]
[TD="class: U2"]U2[/TD]
[TD="class: U3"]U3[/TD]
[TD="class: U4"]U4[/TD]
[TD="class: U5"]U5[/TD]
[TD="class: U"]U[/TD]
[TD="class: K"]K[/TD]
[TD="class: I"]I[/TD]
[TD="class: W"]W[/TD]
[TD="class: X"]X[/TD]
[TD="class: others, width: 40"]Other[/TD]
[TD="class: others, width: 40"]Size[/TD]
Spain2.40.744.1(28)(2.6)7.56.62.16.41.11.41.98.11.86.31.11.41.75.52506
Andalusia7.40.844.3(29.5)4.88.92.32.6111.65.71.26.81.31.63.23.1310
Aragon1.2139.3515.8010.90.801.79.604.21.70.80.87.2119
Asturias0154.15.6901.102.2012.32.27.91.11.102.489
Basques0.30.849(44)(2.8)7.97.61.5610.30.811.71.95.30.61.12.31.8618
Cantabria1.62.537.6(27)193.70.42.50.81.22.910.72.53.72.9000.4242
Catalonia3.10.529.57.571.37.61.32.53.810.13.9101.352.53.180
Northern Spain has more mtDNA U5 than Southern Spain, but Andalusia carries about 7% of mtDNA haplogroups U5 and nearly 10% of y-dna haplogroup I2. It seems unlikely to find zero percent WHG for Central and Southern Spain given the haplogroups listed above. I know some U5 may have come from the Steppe or EHG, but WHG seems a more likely source since Spain was a refugia in the last ice age.
 



Northern Spain has more mtDNA U5 than Southern Spain, but Andalusia carries about 7% of mtDNA haplogroups U5 and nearly 10% of y-dna haplogroup I2. It seems unlikely to find zero percent WHG for Central and Southern Spain given the haplogroups listed above. I know some U5 may have come from the Steppe or EHG, but WHG seems a more likely source since Spain was a refugia in the last ice age.


Sorry, I'm pressed for time, so this will be a cursory response.

Yes, the Haak results giving 0 WHG for both southern Spain and Tuscany stand out. Certainly, amateur analysis shows more WHG than that, in addition to what is in the particular EEF sample used.

Could it be a function of the fact that they used Loschbour?

Just as a general proposition, I also wouldn't use uniparental markers to guess at autosomal component distribution. That can turn out wildly wrong. How much West Eurasian do those West African R1b V88 carry? My father's area is probably 70% R1b, but the amount of steppe there is nowhere near that, probably 30% tops.
 
Thank you so much Mrs. Angela for merging the threads. I had forgotten about this preprint. What struck me about the map of Israel was that although the Jews may have picked up some European DNA, the Anatolian farmers were very different from the Levantine farmers.
 
How does Spain show zero WHG in the Haak chart? They have a significant amount of mtDNA U5 and y-dna I2.

Yes, it's very unlikely that any part of Spain has zero WHG. A bit of WHG might be already inside Early Neolithic (LBK_EN) and maybe there's a little mistake here in Haak's chart.
 
Another point.
A factor that may have contributed to the increase in the percentage of y chromosomes in the farmer populations: As a rule, pre-farmer peoples around the world have a division of labor, women harvest vegetables and men hunt. With agriculture, wild plants practically disappear from cultivated fields while wild animals continue to cross them and even grow there because they have more food. Thus the gatherer women had to move away from the cultivated areas to find wild plants or they were stoned to avoid stealing farmers' crops. The men came closer to these lands because they had more prey and were well received for eliminating the herbivores that devastated the crops, the carnivores that attacked the domestic animals, and the savage ancestors of the domesticated who, when they crossed with them, lowered their desired qualities. Although they could hunt domestic animals the final balance would always be positive for both parties. When the accumulation of diseases and alterations caused by sedentary life (rotten teeth, diabetes, short stature, weaker bones and muscles, ...) led farmer women, to kiss the healthy forest hunter, whose remains have the complete dentition still today (except at least for the areas of pine nut, hazelnut and the like) women gatherers were away. Nor was it necessary for the biological dad of the children born to these encounters to be integrated or even known among the farmers.
 
That's all speculation upon speculation. Plus, it's a bit too much Conan the Barbarian centered, like many of the musings of Eurogenes.) Maybe it happened that way, maybe it didn't. Maybe it was just chance.

I hate to burst your bubble, but the WHG populations were small in number for a reason. The hunter gatherers were always on the verge of extinction because their numbers were so low in comparison to the farmers, and the numbers were low because it takes so much more land to support a hunter than a farmer. They also have no reserves for lean times. Why do you think people so readily adopted agriculture and herding in place of hunting and gathering? As a result, the HGs lived in small, isolated bands, and were sometimes quite inbred. So, if some predator didn't eat them, or a neighbor didn't bash them over the head with a club, maybe they were healthier and maybe they weren't. In the Gamba paper on the Hungarian Neolithic, the HG seems to have been a slave.

It may be that in one of the cyclical farming crashes and the population crashes accompanying them some men carrying yDna I2 got lucky. Maybe in those lean years some WHG with good hunting skills were adopted into the group and their offspring had a better chance of survival. Or maybe, as in the Hungarian Neolithic (Gamba paper), some were enslaved but their descendants got lucky. It happens.

The fact remains that those yDna 12 men in Megalithic tombs, as one example, were 80% farmer. Y dna comes and goes. It's always been like that and it always will be. Men carrying the same yDna can be vastly different from one another in overall genetic similarity.

As to the Haak graphic, I highly doubt there's a mistake in the statistics. The problem may lie with using only one sample, Loschbour, for the comparison. I don't have time to look up the papers, but more than one mentioned that the similarity was more to the Central European WHG.
 
Honorable Mrs. Angela: It seems that what led to the spreading of the neolithic out of the big river valleys (slash-and-burn farming) is that after 3 to 5 years it must chage of place due to the exhaustion of the land, only after 5 to 20 years the fields are productive and with the repetition of the cycle the lands end up forever depleted. We have a dozen different foods, routine and incomplete physical activity, always the same timings and places. May be it's atavism but the separation between town and country seems to me to be schizophernic. HG usually have in their diet 250 to 300 plants and animals ; varied and complete physical activity; harvest, hunt, spend the night in different places and work a lot less hours than we do. I would only abandon this for reasons of force majeure.If the WHG readily adopted agriculture and herding, had increased their populations and we had no anatolian farmers DNA. The same in Asia: andaman like DNA in south asia, andaman like DNA in souteast asia, andaman like DNA in siberia, andaman like DNA in east asia. Where is the "andaman like" y chromossome? only in isolated places (tibet, japan and the tiny andaman islands). If they readily adopted agriculture and herding, had increased their populations and all this places were still 100% andaman like. In Africa they were all 100% Bushmen and the like. Americas, Australia, ... all the same way.
 
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That's all speculation upon speculation. Plus, it's a bit too much Conan the Barbarian centered, like many of the musings of Eurogenes.) Maybe it happened that way, maybe it didn't. Maybe it was just chance.

I hate to burst your bubble, but the WHG populations were small in number for a reason. The hunter gatherers were always on the verge of extinction because their numbers were so low in comparison to the farmers, and the numbers were low because it takes so much more land to support a hunter than a farmer. They also have no reserves for lean times. Why do you think people so readily adopted agriculture and herding in place of hunting and gathering? As a result, the HGs lived in small, isolated bands, and were sometimes quite inbred. So, if some predator didn't eat them, or a neighbor didn't bash them over the head with a club, maybe they were healthier and maybe they weren't. In the Gamba paper on the Hungarian Neolithic, the HG seems to have been a slave.

It may be that in one of the cyclical farming crashes and the population crashes accompanying them some men carrying yDna I2 got lucky. Maybe in those lean years some WHG with good hunting skills were adopted into the group and their offspring had a better chance of survival. Or maybe, as in the Hungarian Neolithic (Gamba paper), some were enslaved but their descendants got lucky. It happens.

The fact remains that those yDna 12 men in Megalithic tombs, as one example, were 80% farmer. Y dna comes and goes. It's always been like that and it always will be. Men carrying the same yDna can be vastly different from one another in overall genetic similarity.

As to the Haak graphic, I highly doubt there's a mistake in the statistics. The problem may lie with using only one sample, Loschbour, for the comparison. I don't have time to look up the papers, but more than one mentioned that the similarity was more to the Central European WHG.

Not to disagree with the bulk of your post about adoption of farming, but I would put a "bemol" on your presentation of the Y-haplo/autosomes (real) dichotomy.
Farmers of Central Europe (LBL) were mostly ANF, and their Y-markers were 'neolithic' too.
IN every region where and when HG's autosomes increased we see an increase of Y-I2a 's haplos, whatever the subclade. Sure the %'s are a bit isconnected, but this has some signification to me. No pure hazard of rare successfull male haplos.
 

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