"I already showed you data. But you don't need data, you need meds."
Quote. Eupedia “In such a scenario, North Africans would have belonged primarily to haplogroup E-M78. E-V13 is more common in Lybia today than anywhere in the Near East, which concords with central North Africa as being a potential source for the European E-V13 (and other subclades of M78). “There is clearly a radiation from the Greece (where E-V13 makes up approximately 30% of the paternal lineages) to the East Mediterranean (where the frequency drops to under 5%).”
A strong argument in favour of E1b1b** crossing DIRECTLY from North Africa ***to southern Italy is that South Italians have more African admixture than people in the Balkans, Greece or Anatolia. This is true of the Northwest African admixture and theEast African (Red sea) admixture. Another argument is that E1b1b has never been found among the dozens of Neolithic Y-DNA samples in the Balkans or Central Europe. The Neoltihic farmers who migrated from the Levant to the Balkans would have brought mostly Southwest Asian admixture and apparently exclusively Y-haplogroup G2a. Many Neolithic sites yielded an occasional "outsider" to the G2a majority, but these were lineages (C1a2, F, I1, I2) that are thought to belong to assimilated (or enslaved) Mesolithic hunter-gatherers. That was very probably the case with E-V13 in Catalonia too. The hypothesis of E1b1b settling in Mediterranean Europe since the Late Paleolithic or Mesolithic would also explain: o 1)...why South Italians and Iberians are remarkably dolichocephalic (long-headed) like North Africans, while North Italians (who are more of Italo-Celtic descent) are quite brachycephalic (broad-headed) like Central Europeans, Eastern Europeans and West Asians. A direct migration from North Africa to South Italy would have resulted not only in higher African admixture in South Italians, but also in a similar morphology. o o o .
o 2)...the almost complete absence of other Paleolithic lineage (notably I2) from southern and central Italy, except in Sardinia, which was presumably not settled by Paleolithic North Africans due to its distance from the nearest coast. It would rather have been settled through Corsica from North Italy by Central European hunter-gatherers (I2a1). The modern Sardinian population distinguishes itself by its higher frequency of non-Mediterranean Mesolithic ancestry (such as WHG) but the nearly complete absence of East African admixture. Sardinia is also the only region of Italy which almost doesn't have any E-V13.
-------------------- Without data you are just another person with an opinion - Deming Posts: 12143 | From: When you have eliminated the impossible, whatever remains, however improbable | Registered: Jun 2007
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posted
Again, you fail repeatedly because you don't have a firm grasp on the general situation. Natufian-like people brought brachycephaly to Europe. If not the sole source, they (i.e. Levantines) were the main contributors of this phenotype in Europe. Mesolithic Europeans mostly had long heads, which they inherited from UP Europeans.
Stop posting non sense and just accept your Ls.
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quote:Originally posted by Xyyman: What’s your argument about E-V13 again? I missed it. Are you saying Natufians are E-V13.
It doesn't matter what my personal views are in regards to E-V13 and the Natufians. What matters is that E-V13 has been found in several Eurasian sites ~5000BC. And E-V13 is younger than E-M2, so if E-V13 can show up this early, E-M2 can, too. But it hasn't, so far.
That's a PITA that YOU have to reconcile given your claims over the years. But you can't. Hence, why you're bending over backwards to make excuses for why it's not found around the Med this early.
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posted
Ha! Ha! Ha! You are such a fraud. It is amazing you have been getting away with the bs you post. The chart shows Natufians are NOT ancestral to Europeans. Further more Lazaridis confirm through SNP Natufians are NOT ancestral to Europeans. So what are you blabbing about E-V13. You are a funny dude. Wasting my time . Quite while you are behind. LOL
-------------------- Without data you are just another person with an opinion - Deming Posts: 12143 | From: When you have eliminated the impossible, whatever remains, however improbable | Registered: Jun 2007
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quote:Originally posted by xyyman: The chart shows Natufians are NOT ancestral to Europeans.
"The chart shows". What chart? The chart you annotated with your attempts at drawing? You're masquerading 'charts' you made up as authoritative sources? Note that your annotations don't even match the original annotations.
quote:Further more Lazaridis confirm through SNP Natufians are NOT ancestral to Europeans.
Prove it. Post the citation.
quote:So what are you blabbing about E-V13.
Translation: damn, I just got my ass handed to me on Hallast et al. I better switch my argument up and start finding new evidence to wish away the evidence of E-V13 in the Avellaner Cave and Carpathian Basin E carriers.
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posted
Lol. LL Cool J is hiding behind his own chart and a small sample size to wish away that E1b1b1a was among the Natufians along with E1b1b1b. I guess he's abandoned spamming his fallacy that E-M2 is too young to be among the Natufians.
LL Cool J has apparently deluded himself into believing that E1b1b1b is not a North African haplogroup, but a hg with Sub Saharan African affinity (you know that's the delusion he tries to massage in his readers' mind with that map).
LL Cool J is working overtime frying his poor brains with his cognitive dissonance-fueled rants. He wants to convince people that Natufians were related to most Sub-Saharan Africans and somehow unrelated to the E-V13 people who share the E-M35 mutation with the Natufian E carriers.
LL Cool J and his shenanigans. He really needs his meds. Instead, he's on a diet of Ls.
posted
to the newbies. Europeans are primarily on the V68/M78 branch of E1b1b. Africans/Natufians are on the Z82Z branch.
But more importantly. Notice, Ethiopians carry ANCESTRAL clades found in the "middle East".
Significance? direction of migration. Lazaridis is blowing smoke about back-migration from the Levant to Ethiopia.
-------------------- Without data you are just another person with an opinion - Deming Posts: 12143 | From: When you have eliminated the impossible, whatever remains, however improbable | Registered: Jun 2007
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
The ignorance displayed in this thread is appalling. It shows iinability to analyze genetic reports and accurately read phylogenies, not to mention lack of subordinating ego to facts.
Trombetta 2015 Fig 1 overtly dates:
E-M2 at 15.3 kya by BEAST or 14.7 kya by rho
E-V13 at 8.1 kya by BEAST or 6.1 kya by rho
Self proclaimed know it all teachers of the masses don't know enough to discern a regional TMRCA from the actual first appearance origin TMRCA.
FIG. 1. — MP tree of haplogroup E obtained with 729 variable positions. Above each branch are reported the haplogroup nomenclature used in the main text (in square brackets) and the number of mutational events defining each branch. Few internal key mutations are indicated as an aid to the reader. Dating estimates are reported in boxes near each node (the values on top and on bottom were obtained with BEAST and rho method, respectively). At the tip of each terminal branch, the haplogroup affiliation of the sequenced sample is reported, using a mutation-based nomenclature according to Trombetta et al. (2011). The ID of the samples from Complete Genomics is also indicated (in parentheses). Colored belts indicate major lineages.
Beniamino Trombetta. Eugenia D’Atanasio Andrea Massaia Marco Ippoliti Alfredo Coppa Francesca Candilio Valentina Coia Gianluca Russo Jean-Michel Dugoujon Pedro Moral Nejat Akar Daniele Sellitto Guido Valesini Andrea Novelletto Rosaria Scozzari Fulvio Cruciani Phylogeographic Refinement and Large Scale Genotyping of Human Y Chromosome Haplogroup E Provide New Insights into the Dispersal of Early Pastoralists in the African Continent
Genome Biol Evol (2015) 7 (7): 1940-1950. doi: 10.1093/gbe/evv118 First published online: June 24, 2015
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posted
I thought the new wife would keep you off the keyboard?
Hallast et al. STR vs SNP dating. Git it! got it?!
You are getting caught-up with the smoke and mirrors also. Even with Thrombetta method of dating E-M2 is NOT 54Kyo as the conman stated. Also as I stated earlier in the thread there are different techniques for dating. That is why there are different dates. But the red herring put out by Swenet was not the initially argument. E-V13 has a North Africa “direct” to Southern Europe migration pattern. The Natufians are NOT ancestral to Europeans. Yes, they migrated FROM SSA but they are also NOT closely related to modern West Africans because as I stated they do NOT carry the E-M2 haplogroup. Which I speculated that E-M2 mutation either did not occur as yet or was NOT established enough to migrate along with E1b1b1b2 to the Levant. As Lazaridis stated obviously the Natufians were pre-Neolithic. Keep in mind E-M2 mutation is of East African origin. East Africans carry E-M2 and the ancestral clades of E-M2 Following the argument.
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quote: Originally posted by xyyman: The Natufians are NOT ancestral to Europeans.
Just stop embarrassing yourself, Samuel L Jackson.
Natufian → PPN → Catal Huyuk & other Anatolian sites → first farming colonists in Greece.
Everybody knows that. Only looney toon clowns like yourself are confused. It's raining Ls in this thread.
quote:Originally posted by Xyyman: Even with Thrombetta method of dating E-M2 is NOT 54Kyo as the conman stated.
LL Cool J still doesn't know the difference between E1b1a1 (~15kya per Trombetta et al) and E1b1a (~45kya per Trombetta et al). And get the date right. I never said 54kya.
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posted
This may be confusing to you but we are talking GENETICS ONLY. Their is no genetic data to support Neolithic of Greece were from the Levant. So, no, everyone does NOT know this.
Oh! and you do know Greeks carry E--V13(E1b1b1a) and NOT the Natufian E1b1b1b. lol! why do I bother?
SMH
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quote:Originally posted by xyyman: This may be confusing to you but we are talking GENETICS ONLY.
It has already been demonstrated that early European farmers crossed the Aegean from Anatolia. Wasn't it you who was bragging about popularizing this paper just days ago?
Doesn't matter what data set you try to run to. If you arbitrarily insist on only accepting population genetics data I have no problem with that. You'll just find more Ls waiting for you in the genetics department.
quote:Originally posted by xyyman: Oh! and you do know Greeks carry E--V13(E1b1b1a) and NOT the Natufian E1b1b1b.
Before that argument can make sense, you first have to prove that the Natufians are exclusively E-Z827. Better yet, prove that it's a normal condition for a population to belong to a single haplogroup branch. Go on. Prove the depths of your ignorance.
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
Once again, repeat until learned
Trombetta 2015 Fig 1 overtly dates:
E-M2 at 15.3 kya by BEAST or 14.7 kya by rho
E-V13 at 8.1 kya by BEAST or 6.1 kya by rho
All else not recognizing the above basic FACT is irrelevant rubbish.
Newbies, and everyone else truly interested , there's no shortcut.
You must link and download Trombetta 2015 to STUDY, - - merely scan read for myopic proofs of your preconceptions - - the report in its particulars, including the dating analysis of past reports.
quote:"This area was like a paradise," says Schmidt, a member of the German Archaeological Institute. Indeed, Gobekli Tepe sits at the northern edge of the Fertile Crescent—an arc of mild climate and arable land from the Persian Gulf to present-day Lebanon, Israel, Jordan and Egypt—and would have attracted hunter-gatherers from Africa and the Levant. And partly because Schmidt has found no evidence that people permanently resided on the summit of Gobekli Tepe itself, he believes this was a place of worship on an unprecedented scale—humanity's first "cathedral on a hill."
posted
Strawman, tactic on your end. Who said a population has ONLY ONE haplogroup? You can see for yourself as presented by Lazaridis that that PPNB and Natufians carry several haplogroups which most are of "recent" Africa origin.
Stop mis-directing.
quote:Originally posted by Swenet:
quote:Originally posted by xyyman: This may be confusing to you but we are talking GENETICS ONLY.
It has already been demonstrated that early European farmers crossed the Aegean from Anatolia. Wasn't it you who was bragging about popularizing this paper just days ago?
Doesn't matter what data set you try to run to. If you arbitrarily insist on only accepting population genetics data I have no problem with that. You'll just find more Ls waiting for you in the genetics department.
quote:Originally posted by xyyman: Oh! and you do know Greeks carry E--V13(E1b1b1a) and NOT the Natufian E1b1b1b.
Before that argument can make sense, you first have to prove that the Natufians are exclusively E-Z827. Better yet, prove that it's a normal condition for a population to belong to a single haplogroup branch. Go on. Prove the depths of your ignorance.
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All else not recognizing the above basic FACT is irrelevant rubbish.
Newbies, and everyone else truly interested , there's no shortcut.
You must link and download Trombetta 2015 to STUDY, - - merely scan read for myopic proofs of your preconceptions - - the report in its particulars, including the dating analysis of past reports.
In that case the best proxy would be:
E-M2 at 15.3 kya by BEAST or 14.7 kya by rho
quote: Cylindrical objects made usually of fired clay but sometimes of stone were found at the Yarmukian Pottery Neolithic sites of Sha‘ar HaGolan and Munhata (first half of the 8th millennium BP) in the Jordan Valley. Similar objects have been reported from other Near Eastern Pottery Neolithic sites. Most scholars have interpreted them as cultic objects in the shape of phalli, while others have referred to them in more general terms as “clay pestles,” “clay rods,” and “cylindrical clay objects.” Re-examination of these artifacts leads us to present a new interpretation of their function and to suggest a reconstruction of their technology and mode of use. We suggest that these objects were components of fire drills and consider them the earliest evidence of a complex technology of fire ignition, which incorporates the cylindrical objects in the role of matches.
[...]
Drilling has been documented as early as the Natufian culture (15,000–11,700 years calBP) through increased numbers of cap stones and drilled stones including beads [26]–[27].
--Naama Goren-Inbar et al.
The Earliest Matches
Received: May 15, 2012; Accepted: July 2, 2012; Published: August 1, 2012
PLoS ONE 7(8): e42213. doi:10.1371/journal.pone.0042213
quote: We report on uniquely preserved 13,700–11,700-y-old grave linings made of flowers, suggesting that such use began much earlier than previously thought.
[...]
The earliest cemeteries (ca. 15,000–11,500 y ago) in the Levant are known from Natufian sites in northern Israel, where dozens of burials reflect a wide range of inhumation practices.
--Dani Nadela,1, Avinoam Daninb, Robert C. Powerc, Arlene M. Rosend, Fanny Bocquentine, Alexander Tsatskina, Danny Rosenberga, Reuven Yeshuruna,f, Lior Weissbroda, Noemi R. Rebollog, Omry Barzilaig, and Elisabetta Boarettog
Earliest floral grave lining from 13,700–11,700-y-old Natufian burials at Raqefet Cave, Mt. Carmel, Israel
quote:The Natufians existed in the Mediterranean region of the Levant 15,000 to 11,500 years ago. Dr. Grosman suggests this grave could point to ideological shifts that took place due to the transition to agriculture in the region at that time.
posted
^^^ useless information, there is no evidence of farming ("neolithic") at Gobekli Tepe at the time it was built
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
Natufians did NOT migrate from Inner Africa. There are absolutely NO Natufian archaeological sites in continental Africa. One Natufian contributing component came from continental Africa and dominates their nrY chromosomes, but Natufians' whole genomes indicate their local component is a major factor that cannot be denied or ignored anymore than any other fact of life.
A lot more's known now since Garrod's original findings. Note the range and extent of Natufian sites. Pay attention to pre and post Natufians as well as Natufufian neighbors.
quote:Originally posted by the lioness,: ^^^ useless information, there is no evidence of farming ("neolithic") at Gobekli Tepe at the time it was built
LOL And who are you actually to dispute any of what was said in that article? LOLThe only thing useless here is you. As you can not even put in data to counteract what was stated in that article. Yes, that's how ridiculous you are. So stick to the topic by putting in data, or don't argue at all.
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quote:Originally posted by xyyman: E-V13 has a North Africa “direct” to Southern Europe migration pattern. The Natufians are NOT ancestral to Europeans.
To make matters worse, xyyman apparently doesn't even known that early neolithic Africans by and large did not specialize in farming. So how could they have possibly popped up in Europe as full-fledged farmers at a time when crops from the levant hadn't even appeared in North Africa?
quote:Strawman, tactic on your end. Who said a population has ONLY ONE haplogroup?
That's EXACTLY what you said when you said "Natufians are on the E1b1b1b branch" and even annotated a map to visualize this bizarre claim. Don't try to backtrack now.
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quote:Originally posted by the lioness,: Ish Gebor at least I reply to your posts "stick to the article" , good one
LOL I did stick to the topic, it flew over your little head. The timing and geographic location is coherent with the topic. But you probably did not know that. lol
quote:From this perch 1,000 feet above the valley, we can see to the horizon in nearly every direction. Schmidt, 53, asks me to imagine what the landscape would have looked like 11,000 years ago, before centuries of intensive farming and settlement turned it into the nearly featureless brown expanse it is today.
quote:Originally posted by xyyman: [Q] yes! we are at the end. Lazaridis made clear. A possible pre-Neolithic AFRICAN origin of the Natufians.
The End.
[QUOTE]Originally posted by xyyman: [Q] There you have it!!!! Natufians were of African origin. Which contributed to Basal Eurasian. Europeans are as much as 80% Africans. didn't I tell you that. Europeans are a subset of Africans.
It is all in this one paragraph.
Persian Neolithic is also African origin.
DNATribes was correct. These African neolithics brought the technology up from the Harrapan Valley to Iberia including Greece and Sardinia.
========== Quote:
"Our finding that at least Natufians had Y-chromosomes of ****African origin ****(Supplementary Information, section 6) suggested to us ***initially *****that gene flow from Africa (which did not experience Neanderthal admixture) may have contributed Basal Eurasian ancestry into the ancient Near East, However, Natufians do not have less Neanderthal ancestry (Fig. S5.1) or more Basal Eurasian ancestry (Supplementary Information, section 4) than the Neolithic of Iran, and so do not appear to be exceptional in either respect within the context of the ancient Near East. The presence of Basal Eurasian ancestry not only in the Epipaleolithic Natufians from the Levant, but also of the Upper Paleolithic/Mesolithic of Georgia8 suggest that while this type of ancestry first appears in Europe with the Early Neolithic7, it was already pervasive in the Near East before the advent of the Neolithic. Future studies of human remains from the Near East may determine (i) how much earlier the Basal Eurasians were present there, (ii) whether they represent a population of African origin or one that lived in Eurasia but did not experience the Neanderthal admixture of other Eurasians, and (iii) when the dilution of Neanderthal admixture first took place." [/Q]
[/][/TE]
quote:Originally posted by Tukuler: [Q] Natufians did NOT migrate from Inner Africa. There are absolutely NO Natufian archaeological sites in continental Africa. One Natufian contributing component came from continental Africa and dominates their nrY chromosomes, but Natufians' whole genomes indicate their local component is a major factor that cannot be denied or ignored anymore than any other fact of life.
A lot more's known now since Garrod's original findings. Note the range and extent of Natufian sites. Pay attention to pre and post Natufians as well as Natufufian neighbors. [/QB]
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posted
xyyman; Europeans are depigmented Africans but the Natufians were brother men with elite farming skilz
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
C'mon man, ain't I already hipped you the Y hg tells but a miniscule story compared to the WHOLE fucking GENOME. Was your eats all waxed or didn't you hear
Get it? Got it!
I could drudge up old mistaken notions I had about Natufians when I arrived at ES just to show that I finally shut up listened and learned from the now long gone vets.
You need Natufian specialists writers not geneticist if you want to learn then go teach what you can't get in a little paragraph.
You gotta do more than call Bullshit. Get off the net and visit a library sometime to do an interlibrary loan or spelunk a special collection.
A freebie for you, Africanists know what Inner Africa is, so why don't you. Even Keita knew Inner Africa from Coastal Africa.
North Coastal Africa vs Inner Africa - Cape Alguhas to the pre-Sahara
.
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
@ Ish
That map? Man, that's the s h I t , man.
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You are preaching to the converted about phenotype and genotype. What you like many others fail to realize is that there was never no “isolation of population”. Rosenberg et al. That is why I am way ahead than the others. Once you accept the fact and understand it is a continuum and genetic surfing explain everything. You will be ahead of the game.
quote; "C'mon man, ain't I already hipped you the Y hg tells but a miniscule story compared to the WHOLE fucking GENOME."
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“Love you some books”. These books are being replaced. You know that? Don’t you?. Again, what is “inner Africa”. What proof do you have inner Africa was occupied and by what ethnic group? Dogma.
Signed the Africanist.
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
Correct
Genetics don'tie.
The whole genome and nothing but the whole genome so help me Molecular Geneticist
U still the same U na wanna learn U wanma count coup and apply the Ostrich Theory Yr head hidden in the sand Yr asz out in the air
posted
LL Cool J's highly specialized non-farming "E-V13 farmers" who thought the Pelasgians how to domesticate and cultivate crops.
quote:For Africa, these patterns do not apply. There, the Neolithic Revolution followed a different path –the main differences are that pottery was present in Africa around 2000 years prior to cultivated cereals (Haaland 2007: 165; Linseele 2010: 43) and that its invention was independent to food production (Hassan1998: 89), but also, that cereal cultivation developed after animal domestication (Marshall and Hildebrand 2002: 101).
All these acrobatics and shenanigans because LLman is salty about the absence of E-M2 in Eurasian aDNA so far. SMH.
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posted
There will NOT any E-M2 prior to 4000BC WITHIN West Africa!!! It is a new mutation.
There will not be E-M2 OUTSIDE Africa prior to 4000BC. It is a new mutation
"Nihga please I wz into this before u knew there wz such a thing as ES"
I never said otherwise. I learnt a lot from you. Sometimes the pupil become the teacher.
-------------------- Without data you are just another person with an opinion - Deming Posts: 12143 | From: When you have eliminated the impossible, whatever remains, however improbable | Registered: Jun 2007
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
In this case the pupil became a dunce.
Anyone wants ta swear by you, in the face of molecular biologists reporting E-M2's greater than 14kya coalescence, deserves the morass they'll find them selves quagmired in.
Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
My last post was truncated. The part about your two major breakthroughs re no SW Europe refugium pour out in NW Africa and checking mummy STRs on our own were great teachings I imbibed from you.
As I say in real life to my students when they ask who are my teachers; I reply ALL OF THEM
The more and more I learn the less I'm sure I know.
1. Neanderthal were black skinned 2. The Bantu expansion probably never occurred 3. and many others
agreed. I followed your postings to get me on the right track. Along with a few others like Rasol, Lion(not lioness), Jari, etc. I always had doubts about DJ and Ausar based upon their tone-nation.
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Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
Well I was only listing your original stuff that informed me. That the baNtu expansion was lingocultual not demic is something I posted more than once before your day.
I llearned it from the coauthor of one of UNESCO's history volumes whose fellow. coauthor had the editor disavow sny connection to his name.
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Early farmers from across Europe directly descended from Neolithic Aegeans Zuzana Hofmanováa, et al 2016
[excerpts]
Significance
One of the most enduring and widely debated questions in prehistoric archaeology concerns the origins of Europe’s earliest farmers: Were they the descendants of local hunter-gatherers, or did they migrate from southwestern Asia, where farming began? We recover genome-wide DNA sequences from early farmers on both the European and Asian sides of the Aegean to reveal an unbroken chain of ancestry leading from central and southwestern Europe back to Greece and northwestern Anatolia. Our study provides the coup de grâce to the notion that farming spread into and across Europe via the dissemination of ideas but without, or with only a limited, migration of people.
Abstract
Farming and sedentism first appeared in southwestern Asia during the early Holocene and later spread to neighboring regions, including Europe, along multiple dispersal routes. Conspicuous uncertainties remain about the relative roles of migration, cultural diffusion, and admixture with local foragers in the early Neolithization of Europe. Here we present paleogenomic data for five Neolithic individuals from northern Greece and northwestern Turkey spanning the time and region of the earliest spread of farming into Europe. We use a novel approach to recalibrate raw reads and call genotypes from ancient DNA and observe striking genetic similarity both among Aegean early farmers and with those from across Europe. Our study demonstrates a direct genetic link between Mediterranean and Central European early farmers and those of Greece and Anatolia, extending the European Neolithic migratory chain all the way back to southwestern Asia.
It is well established that farming was introduced to Europe from Anatolia, but the extent to which its spread was mediated by demic expansion of Anatolian farmers, or by the transmission of farming technologies and lifeways to indigenous hunter-gatherers without a major concomitant migration of people, has been the subject of considerable debate. Paleogenetic studies [1⇓⇓–4] of late hunter-gatherers [HG] and early farmers indicate a dominant role for migration in the transition to farming in central and northern Europe, with evidence of only limited hunter-gatherer admixture into early Neolithic populations, but increasing toward the late Neolithic. However, the exact origin of central and western Europe’s early farmers in the Balkans, Greece, or Anatolia remains an open question.
Recent radiocarbon dating indicates that by 6,600–6,500 calibrated [cal] BCE sedentary farming communities were established in northwestern Anatolia at sites such as Barcın, Menteşe, and Aktopraklık C and in coastal western Anatolia at sites such as Çukuriçi and Ulucak, but did not expand north or west of the Aegean for another several hundred years [5]. All these sites show material culture affinities with the central and southwestern Anatolian Neolithic [6].
Early Greek Neolithic sites, such as the Franchthi Cave in the Peloponnese, Knossos in Crete, and Mauropigi, Paliambela, and Revenia in northern Greece date to a similar period [7⇓–9]. The distribution of obsidian from the Cycladic islands, as well as similarities in material culture, suggest extensive interactions since the Mesolithic and a coeval Neolithic on both sides of the Aegean [8]. Although it has been argued that in situ Aegean Mesolithic hunter-gatherers played a major role in the “Neolithization” of Greece [7], the presence of domesticated forms of plants and animals indicates nonlocal Neolithic dispersals into the area.
We present five ancient genomes from both, the European and Asian sides of the northern Aegean [Fig. 1]; despite their origin from nontemperate regions, three of them were sequenced to relatively high coverage [∼2–7×], enabling diploid calls using a novel SNP calling method that accurately accounts for postmortem damage [SI Appendix, SI5. Genotype Calling for Ancient DNA]. Two of the higher-coverage genomes are from Barcın, south of the Marmara Sea in Turkey, one of the earliest Neolithic sites in northwestern Anatolia [individuals Bar8 and Bar31]. On the European side of the Aegean, one genome is from the early Neolithic site of Revenia [Rev5], and the remaining two are from the late and final Neolithic sites of Paliambela [Pal7] and Kleitos [Klei10], dating to ∼2,000 y later [Table 1]. Estimates of mitochondrial contamination were low [0.006–1.772% for shotgun data] [Table 1; SI Appendix, SI4. Analysis of Uniparental Markers and X Chromosome Contamination Estimates.]. We found unprecedented deamination rates of up to 56% in petrous bone samples, indicating a prehistoric origin for our sequence data from nontemperate environments [SI Appendix, Table S5].
Uniparental Genetic Systems
The mtDNA haplogroups of all five Neolithic individuals are typical of those found in central European Neolithic farmers and modern Europeans, but not in European Mesolithic hunter-gatherers [1]. Likewise, the Y-chromosomes of the two male individuals belong to haplogroup G2a2, which has been observed in European Neolithic farmers [3, 10]; in Ötzi, the Tyrolean Iceman [11]; and in modern western and southwestern Eurasian populations, but not in any pre-Neolithic European hunter-gatherers [12]. The mitochondrial haplogroups of two additional less well-preserved Greek Mesolithic individuals [Theo1, Theo5; SI Appendix, Table S6] belong to lineages observed in Neolithic farmers from across Europe; consistent with Aegean Neolithic populations, unlike central European Neolithic populations, being the direct descendants of the preceding Mesolithic peoples who inhabited broadly the same region. However, we caution against over-interpretation of the Aegean Mesolithic mtDNA data; additional genome-level data will be required to identify the Mesolithic source population[s] of the early Aegean farmers.
Functional Variation Sequences in and around genes underlying the phenotypes hypothesized to have undergone positive selection in Europeans indicate that the Neolithic Aegeans were unlikely to have been lactase persistent but carried derived SLC24A5 rs1426654 and SLC45A2 rs16891982 alleles associated with reduced skin pigmentation. Because our Aegean samples predate the period when the rs4988235 T-allele associated with lactase persistence in Eurasia reached an appreciable frequency in Europe, around 4 kya [12⇓–14], and because this allele remains at relatively low frequencies [<0.15] in modern Greek, Turkish, and Sardinian populations [15], this observation is unsurprising. However, despite their relatively low latitude, four of the Aegean individuals are homozygous for the derived rs1426654 T-allele in the SLC24A5 gene, and four carry at least one copy of the derived rs16891982 G-allele in the SLC45A2 gene. This suggests that these reduced-pigmentation–associated alleles were at appreciable frequency in Neolithic Aegeans and that skin depigmentation was not solely a high-latitude phenomenon [SI Appendix, SI12. Functional Markers]. The derived rs12913832 G-allele in the HERC2 domain of the OCA2 gene was heterozygous in one individual [Klei10], but all other Aegeans for whom the allelic state at this locus could be determined were homozygous for the ancestral allele, indicating a lack of iris depigmentation in these individuals.
Examination of several SNPs in the TCF7L2 gene region indicates that the two Neolithic Anatolian individuals, Bar8 and Bar31, are likely to have carried at least one copy of a haplotype conferring reduced susceptibility to type 2 diabetes [T2D]; the Klei10 and Rev5 individuals also carry a tag allele associated with this haplotype. Consistent with these observations, it has been previously estimated that this T2D-protective haplotype, which shows evidence for selection in Europeans, East Asians, and West Africans, originated ∼11,900 y ago in Europe [16].
A number of loci associated with inflammatory disease displayed the derived alleles, including rs2188962 C > T in the SLC22A5/IRF1 region, associated with Crohn’s disease; rs3184504 C > T in the SH2B3/ATXN2 region, associated with rheumatoid arthritis, celiac disease, and type 1 diabetes; and rs6822844 G > T in the IL2/IL21 region, associated with rheumatoid arthritis, celiac disease, and ulcerative colitis. Interestingly, we observe derived states for six of eight loci in a protein–protein interaction network inferred to have undergone concerted positive selection 2.6–1.2 kya in Europeans [17], suggesting that any recent selection on these loci acted on standing variation present at already appreciable frequency [SI Appendix, SI12. Functional Markers]. Fig. 2. PCA of modern reference populations [18, 19] and projected ancient individuals. The Greek and Anatolian samples reported here cluster tightly with other European farmers close to modern-day Sardinians; however, they are clearly distinct from previously published Caucasian hunter-gatherers [20]. This excludes the latter as a potential ancestral source population for early European farmers and suggests a strong genetic structure in hunter-gatherers of Southwest Asia. Central and East European [C./E. European], South European [South Eur.]. Ancient DNA data: Pleistocene hunter-gatherer [Plei. HG] [20, 21, 22], Holocene hunter-gatherer [Holocene HG] [2, 4, 13, 20, 23], Neolithic [2, 4, 12, 13, 24], Late Neolithic/Chalcolithic/Copper Age [LN/Chalc./CA] [13, 25], and Bronze Age [13]. Ancient samples are abbreviated consistently using the nomenclature “site-country code-culture”; see SI Appendix, Table S14 and Dataset S1 for more information. A 3D PCA plot can be viewed as a 3D figure [https://figshare.com/articles/Hofmanova_et_al_3D_figure_S4/3188767].
To examine this clustering of Early Neolithic farmers in more detail, we calculated outgroup f3 statistics [26] of the form f3 [‡Khomani; TEST, Greek/Anatolian], where TEST is one of the available ancient European genomes [SI Appendix, SI7. Using f-statistics to Infer Genetic Relatedness and Admixture Amongst Ancient and Contemporary Populations and Figs. S8–S10; Dataset S2]; ‡Khomani San were selected as an outgroup as they are considered to be the most genetically diverged extant human population. Consistent with their PCA clustering, the northern Aegean genomes share high levels of genetic drift with each other and with all other previously characterized European Neolithic genomes, including early Neolithic from northern Spain, Hungary, and central Europe. Given the archaeological context of the different samples, the most parsimonious explanation for this shared drift is migration of early European farmers from the northern Aegean into and across Europe [12].
To better characterize this inferred migration, we modeled ancient and modern genomes as mixtures of DNA from other ancient and/or modern genomes, a flexible approach that characterizes the amount of ancestry sharing among multiple groups simultaneously [18, 27] [Fig. 3; SI Appendix, SI10. Comparing Allele Frequency Patterns Among Samples Using a Mixture Model]. Briefly, we first represented each ancient or modern “target” group by the [weighted] number of alleles that they share in common with individuals from a fixed set of sampled populations [i.e., the “unlinked” approach described in ref. 27], which we refer to as the “allele-matching profile” for that target group. To cope with issues such as unequal sample sizes, we then used a linear model [28] to fit the allele-matching profile of the target group as a mixture of that of other sampled groups. Sampled groups that contribute most to this mixture indicate a high degree of shared ancestry with the target group relative to other groups. Under this framework the oldest Anatolian genome [Bar31] was inferred to contribute the highest amount of genetic ancestry [39–53%] to the Early Neolithic genomes from Hungary [13] and Germany [2] compared with any other ancient or modern samples, with the next highest contributors being other ancient Aegean genomes [Klei10, Pal7, Bar8] [SI Appendix, Figs. S23, S24, and S29]. This pattern is not symmetric in that we infer smaller contributions from the German [<26%] and Hungarian [<43%] Neolithic genomes to any of the Anatolian or Greek ancient genomes. Furthermore, in this analysis modern samples from Europe and surrounding regions are inferred to be relatively more genetically related to the Aegean Neolithic genomes than to the Neolithic genomes from Germany and Hungary [Fig. 3; SI Appendix, SI10. Comparing Allele Frequency Patterns Among Samples Using a Mixture Model]. These patterns are indicative of founder effects [29] in the German and possibly Hungarian Neolithic samples from a source that appears to be most genetically similar to the Aegean Neolithic samples [specifically, Bar31] and that distinguishes them from the ancestors of modern groups. Consistent with this, we found fewer short runs of homozygosity [ROH] [between 1 and 2 Mb] in our high-coverage Anatolian sample [Bar8] than in Early Neolithic genomes from Germany and Hungary [SI Appendix, SI11. Runs of Homozygosity and Fig. S31]. However, it is not possible to infer a direction for dispersal within the Aegean with statistical confidence because both the Greek and Anatolian genomes copy from each other to a similar extent. We therefore see the origins of European farmers equally well represented by Early Neolithic Greek and northwestern Anatolian genomes.
Ongoing gene flow into and across the Aegean is also indicated in the genome of a Chalcolithic individual from Kumtepe [Kum6 [25]], a site geographically close to Barcın but dating to ∼1,600 y later. Although archaeological evidence indicates a cultural break in many Aegean and West Anatolian settlements around 5,700/5,600 cal BCE [i.e., spanning this 1,600-y period [30]], Kum6 shows affinities to the Barcın genomes in “outgroup” f3-statistics in the form f3 [‡Khomani; TEST, Greek/Anatolian]. The shared drift between Kum6 and both the early and late Neolithic Aegeans is similar in extent to the drift that Aegeans share with one another. However, f4 statistics of the form f4 [Aegean, Kum6, Early farmer, ‡Khomani] were often significantly positive [SI Appendix, Table S22; Dataset S2], suggesting that European Neolithic farmers [namely, Linearbandkeramik [LBK], Starcevo, and Early Hungarian Neolithic farmers] share some ancestry with early Neolithic Aegeans that is absent in Kum6. This is consistent with population structure in the Early Neolithic Aegean or with Kum6 being sampled from a population that differentiated from early Neolithic Aegeans after they expanded into the rest of Europe. Accordingly, compared with Barcın, Kum6 shares unique drift with the Late Neolithic genomes from Greece [Klei10 and Pal7], consistent with ongoing gene flow across the Aegean during the fifth millennium and with archaeological evidence demonstrating similarities in Kumtepe ceramic types with the Greek Late Neolithic [31]. Finally, the Kum6, Klei10, and Pal7 genomes show signals of Caucasus hunter-gatherer [20] admixture that is absent in the Barcın genomes, suggesting post early Neolithic gene flow into the Aegean from the east.
It is widely believed that farming spread into Europe along both Mediterranean and central European routes, but the extent to which this process involved multiple dispersals from the Aegean has long been a matter of debate [32]. We calculated f4 statistics to examine whether the Aegean Neolithic farmers shared drift with genomes from the Spanish Epicardial site Els Trocs in the Pyrenees [3, 12] that is distinct from that shared with Early Neolithic genomes from Germany and Hungary. In a test of the form f4 [Germany/Hungary EN, Spain EN, Aegean, ‡Khomani], we infer significant unique drift among Neolithic Aegeans [not significantly in Bar8] and Early Neolithic Spain to the exclusion of Hungarian and German Neolithic genomes [SI Appendix, Table S21]. The best explanation for this observation is that migration to southwestern Europe started in the Aegean but was independent from the movement to Germany via Hungary. This is also supported by other genetic inferences [24] and archaeological evidence [33]. An alternative scenario is a very rapid colonization along a single route with subsequent gene flow back to Greece from Spain. Potentially, preexisting hunter-gatherer networks along the western Mediterranean could have produced a similar pattern, but this is not supported by archaeological data. Interestingly, Ötzi the Tyrolean Iceman [11] shows unique shared drift with Aegeans to the exclusion of Hungarian Early Neolithic farmers and Late and Post Neolithic European genomes and feasibly represents a relict of Early Neolithic Aegeans [SI Appendix, SI7. Using f-statistics to Infer Genetic Relatedness and Admixture Amongst Ancient and Contemporary Populations and Table S18].
Previous Section Next Section Hunter-Gatherer Admixture Given that the Aegean is the likely origin of European Neolithic farmers, we used Bar8 and Bar31 as putative sources to assess the extent of hunter-gatherer admixture in European farmers through the Neolithic. f4 statistics of the form f4 [Neolithic farmer, Anatolian, HG, ‡Khomani] indicated small but significant amounts of hunter-gatherer admixture into both Spanish and Hungarian early farmer genomes, and interestingly, the Early Neolithic Greek genome. Our mixture modeling analysis also inferred a small genetic contribution from the Loschbour hunter-gatherer genome [3–9%] to each of the Early Neolithic Hungarian and German genomes, but evidence of a smaller contribution to any Aegean genomes [0–6%]. These results suggest that mixing between migrating farmers and local hunter-gatherers occurred sporadically at low levels throughout the continent even in the earliest stages of the Neolithic. However, consistent with previous findings [3], both f4 statistics and ADMIXTURE analysis indicate a substantial increase in hunter-gatherer ancestry transitioning into the Middle Neolithic across Europe, whereas Late Neolithic farmers also demonstrate a considerable input of ancestry from steppe populations [SI Appendix, SI8. Proportions of Ancestral Clusters in Neolithic Populations of Europe and Fig. S32].
Previous Section Next Section Relation to Modern Populations Most of the modern Anatolian and Aegean populations do not appear to be the direct descendants of Neolithic peoples from the same region. Indeed, our mixture model comparison of the Aegean genomes to >200 modern groups [2] indicates low affinity between the two Anatolian Neolithic genomes and six of eight modern Turkish samples; the other two were sampled near the Aegean Sea at a location close to the site of the Neolithic genomes. Furthermore, when we form each Anatolian Neolithic genome as a mixture of all modern groups, we infer no contributions from groups in southeastern Anatolia and the Levant, where the earliest Neolithic sites are found [SI Appendix, Figs. S22 and S30 and Table S30; Dataset S3]. Similarly, comparison of allele sharing between ancient and modern genomes to those expected under population continuity indicates Neolithic-to-modern discontinuity in Greece and western Anatolia, unless ancestral populations were unrealistically small [SI Appendix, SI9. Population Continuity]. Instead, our mixing analysis shows that each Aegean Neolithic genome closely corresponds to modern Mediterraneans [>68% contributions from southern Europe] and in particular to Sardinians [>25%], as also seen in the PCA and outgroup f3 statistics with few substantial contributions from elsewhere. Modern groups matching the Neolithics—mostly from the Mediterranean and North Africa—strikingly match more to Bar8 from northwestern Anatolia than to the LBK genome from Stuttgart in Germany, indicating that the LBK genome experienced processes such as drift and admixture that were independent from the Mediterranean expansion route, consistent with the dual expansion model.
Previous Section Next Section Concluding Remarks Over the past 7 years, ancient DNA studies have transformed our understanding of the European Neolithic transition [1⇓⇓–4, 12, 13], demonstrating a crucial role for migration in central and southwestern Europe. Our results further advance this transformative understanding by extending the unbroken trail of ancestry and migration all of the way back to southwestern Asia.
The high levels of shared drift between Aegean and all available Early Neolithic genomes in Europe, together with the inferred unique drift between Neolithic Aegeans and Early Neolithic genomes from Northern Spain to the exclusion of Early Neolithic genomes from central Europe, indicate that Aegean Neolithic populations can be considered the root for all early European farmers and that at least two independent colonization routes were followed.
A key remaining question is whether this unbroken trail of ancestry and migration extends all the way back to southeastern Anatolia and the Fertile Crescent, where the earliest Neolithic sites in the world are found. Regardless of whether the Aegean early farmers ultimately descended from western or central Anatolian, or even Levantine hunter-gatherers, the differences between the ancient genomes presented here and those from the Caucasus [20] indicate that there was considerable structuring of forager populations in southwestern Asia before the transition to farming. The dissimilarity and lack of continuity of the Early Neolithic Aegean genomes to most modern Turkish and Levantine populations, in contrast to those of early central and southwestern European farmers and modern Mediterraneans, is best explained by subsequent gene flow into Anatolia from still unknown sources.
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I like to put these things up with more text, unadulterated charts, free of comment and with the links
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