November 26, 2015

Neolithic farmers from Greece and Anatolia

A couple of new papers appeared this week. First, an article in Nature on natural selection in ancient Europe includes a sample of Anatolian Neolithic farmers and concludes that the European Neolithic farmers were descended from them with a bit of extra European hunter-gatherer admixture. Second, a new preprint on the bioRxiv includes Neolithic samples from northern Greece and finds that they too resemble the Anatolian and European farmers. I think it is time to declare the problem of "Neolithization of Europe" done. It took less than 4 years to solve it with ancient DNA. Here is a (non-exhaustive) list of papers in historical review:

Nature (2015) doi:10.1038/nature16152

Genome-wide patterns of selection in 230 ancient Eurasians

Iain Mathieson et al.

Ancient DNA makes it possible to observe natural selection directly by analysing samples from populations before, during and after adaptation events. Here we report a genome-wide scan for selection using ancient DNA, capitalizing on the largest ancient DNA data set yet assembled: 230 West Eurasians who lived between 6500 and 300 BC, including 163 with newly reported data. The new samples include, to our knowledge, the first genome-wide ancient DNA from Anatolian Neolithic farmers, whose genetic material we obtained by extracting from petrous bones, and who we show were members of the population that was the source of Europe’s first farmers. We also report a transect of the steppe region in Samara between 5600 and 300 BC, which allows us to identify admixture into the steppe from at least two external sources. We detect selection at loci associated with diet, pigmentation and immunity, and two independent episodes of selection on height.



Early farmers from across Europe directly descended from Neolithic Aegeans

Zuzana Hofmanová, Susanne Kreutzer et al.

Farming and sedentism first appear in southwest 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 Neolithisation of Europe. Here we present paleogenomic data for five Neolithic individuals from northwestern Turkey and northern Greece, spanning the time and region of the earliest spread of farming into Europe. We 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.



Africa Gómez said...

Brilliant summary of the swift advances in this field!

Krefter said...


What people do you think came into Greece after 4000 BC? It doesn't seem most were Yamnaya-types. I'm thinking unknown peoples from West Asia migrated there after 4000 BC and mixed with EEFs. I suspect they brought most J2(CHG?) and E1b-V13. R1b-Z2103 and R1b-M269(xL23) in Greece could also be of CHG or just West Asian origin.

The genetic divide between Turkey and Greece is surprising. I'll investigate it more with genome blogger ADMIXTURE, formals stats, and PCAs. But I'm pretty sure it is large though.

So, another question is why is Greece and Turkey today differnt, while in the Neolithic both were EEF? It must be because differnt people went to each locations and or contributed differnt percentages to each location.

Annie Mouse said...

Jury still out for me.

One Anatolian does not make a source. We dont have any southern Europeans from that time period or earlier. No Italians, No Mediterranean Spaniards. Also No North Africans or folk from the Lebanon etc.

My working hypothesis is this genetic population is the folk floating about around the Med and their cousins. All if this is consistent so far.

Pneumatikon said...

This survey is good genome-wide, but it muddies the water dramatically for us J2a's. These Neolithic farmers they're talking about are largely G2a, and they had a very different fate from us. So I suspect we've got a Hunter-Gatherer snapshot of us before we moved into Central and Southern Anatolia. But after that we look a lot like those G2a men; whose samples come from NW Turkey.

That indicates to me we were a very small group. Makes sense. We were a tiny group when we built the Roman Empire, too.

As for the Sintashta - I have a strong suspicion the "Neolithic Farmer" mix is J2a Balkan descendants already living there, and not men directly from the Balkans after the Kurgan invasion. We were the first to move into the Copper Age, and maps show Copper technology was is use there at the same time it was popping up in the Balkans and Central Anatolia.

The spread along the bottom from Anatolia through Chalcolithic Spain - but NOT including the Remedello and the LBK - matches us beautifully. We probably had a trade network stretching from Spain through the Cyclades, and before my particular J2a line moved to Crete our center of gravity was probably Malta; where we built the first free-standing Megalithic structures. Flanking that site is the Los Millares site in Spain and a splendid settlement on Andros. Both are surrounded stone walls. Both are clearly Anatolians. Nobody else is doing that. The Remedello/LBK complex is probably the G2a line.

I'll buy this article as soon as I get paid next Friday and review it more thoroughly.

Nirjhar007 said...

Will be crucial also to see which Y-DNA those mycenaean folks carry, I expect them to have loads of J2.

Nirjhar007 said...

What people do you think came into Greece after 4000 BC?
I certainly think by mid third millennium bc (Around Liniear A ) , The IE surely arrived in Greece.
With J2 i expect some R1b and even R1a-M417..

mooreisbetter said...

I agree this generally shows a pattern, but it's very easy with aDNA to get caught up in the theory du jour, and forget how much such theories have evolved over time. It's also very easy to use terms such as "Neolithic," "Mesolithic," etc., and forget that they refer to material culture, and not absolute dates.

This post is a bit dated, but it maps the Haplogroups of aDNA based on TIME.

As you can see, the picture is significantly muddy. Especially with G2.

Roy King said...

Despite the fact that I was a big proponent of J2 representing first farmers, I now stand corrected, it was G2a and H2-M282 and T. J2a likely adopted farming and dispersed to Europe from the Chalcolithic Era through the Late Bronze Age. Did J2a bring a wine culture to the Levant, Crete, and Greece? Did the Minoans, Cycladic people, EBA Trojans and LBA Mycenaeans have small colonies in Italy and Spain? These questions linger.

Gaspar said...

In LBK_EN periods in central Europe and the Balkans we have so far
about dozen G2a, 2 x T1a, 6 or so I2a and 1 xH2 . ( remedello was I2a and well as the new Swiss finding Bichon )

The only place where these haplogroups met in ancient times was in modern azeri lands.
IMO, this is the place to looks.

Is it not the case that Mycenaeans are completely different to the Doric race in Greece today?

Andrew Lancaster said...

Great summary. I see that some cases might indeed be closed, of course, but I am not certain about the last bit: that the first wave of "Sardinian like" farmers came only on a route which went through Turkey, then Greece. Archaeologists have long thought that the first wave into Greece might have largely come on an island hopping route, and so the next thing we need is ancient DNA from the Levant, which may well turn out to also look Sardinian? Being more controversial, is there even a chance that farmers in North Africa also looked Sardinian, allowing for a Tunisian route?

Probably, with more ancient DNA, we can identify more sub-routes from this period. Archaeologists already felt there were at least two (LBK and Cardial).

Slumbery said...


We do not know if they met in modern Azeri lands, we do not have samples from there. If I have to guess I would say yDNA haplogroup J was probably present in modern Azeri lands since very ancient times and its absence in Neolithic European and Anatolian samples, together with the absence of CaucasusHG-related autosomal ancestry, makes this theory rather implausible. Even if the migration had sources farther that Western Anatolia (quite possible), a South-Eastern direction (Levant) is much more likely.

Modern day Azerbaijan is a more likely source for the root of the second wave.

andrew said...

@ Pneumatikon Count me out on the "we" language.

@ Krefter Good questions. "What people do you think came into Greece after 4000 BC? It doesn't seem most were Yamnaya-types. I'm thinking unknown peoples from West Asia migrated there after 4000 BC and mixed with EEFs. I suspect they brought most J2(CHG?) and E1b-V13. R1b-Z2103 and R1b-M269(xL23) in Greece could also be of CHG or just West Asian origin."

My working hypothesis is that immediately pre-Mycenaean people of Greece were non-Indo-European Hattic or Hurrian language speakers from Anatolia, at least in most of the Aegean Islands if not on the mainland. I think that J2 is probably from the Caucasus, although the Iranian highlands are also a plausible source.

Nirjhar007 is almost surely wrong in concluding that Linear A, the language of the Minoans, is Indo-European. This is an idea firmly rejected by the linguistic evidence. For example, phonetic translations of Minoan healing spells from Egyptian sources are quite similar to Hattic linguistically, but very unlike any Indo-European language.

I suspect that E1b-V13 derives from Egypt, perhaps in the prehistoric era, or perhaps even in an event recorded in legendary history, the tale of the Danaids fleeing from Egypt seek asylum from King Pelasgus of Argos. Notably, Y-DNA E seems to be absent in ancient Vinca culture DNA.

I suspect that R1b in Greece has the same origins as it does in Western Europe, but with more haplotype diversity due to serial founder effects.

"So, another question is why is Greece and Turkey today differnt, while in the Neolithic both were EEF? It must be because differnt people went to each locations and or contributed differnt percentages to each location."

Lots of layers there. There were Mesopotamian trade colonies in Anatolia and not Greece. The Hittite empire included Turkey but not Greece. The Ottomans had more demic impact in Turkey than in Greece. The Turks had a pervasive impact on Anatolia and almost none on Greece. And, in modern history, there were a Greece-Turkish population exchange and the Armenian genocide that probably remove genetic populations that were intermediate between modern Turkey and modern Greece. Also, Turkey probably had more Caucasian influence.

@ Gaspar

"Is it not the case that Mycenaeans are completely different to the Doric race in Greece today?"

The Mycenaeans are very likely the first Indo-Europeans in Greece and ancestral to modern Greece, albeit with substrate influences and various impacts of Bronze Age and later migrations. There probably is substantial discontinuity with non-Indo-European the immediate predecessors of the Mycenaeans and the Mycenaean Greeks and their descendants.

mooreisbetter said...


[using one's best bullhorn voice]

"Step away from the Kool-Aid. Repeat after me: You are NOT your Haplogroup. Repeat: you are not your Haplogroup!"

The changing nature of populations over time should make this extremely obvious, but apparently it didn't. The populations that were initially I2, J2, R1 -- whatever -- bear little resemblance to the purported "pure" host population.

In other words -- even if the sex-selective wild fantasies popular in the R1b echo chamber otherwise known as Anthrogenica were true -- and even if you had a mass invasion of randy males of R1b, J2, or whomever the theorists (who just HAPPEN to be R1b or J2...) postulate -- then that blood is so diluted over time.

You understand that concept surely that just 6 generations back, you have 64 relatives? The amount of DNA you share with that direct male ancestor is just 1.56%. I strongly suggest checking out the following resources

So unless every male in your pedigree is J2 (and I am unaware of any modern population approaching those saturation levels, except extreme isolates), then you have JUST AS MUCH DNA from all of the other Hgs that make the mosaic of modern populations, especially in Europe.

I'm sorry if I seem harsh, but the pseudoscience on these boards has got to stop. This is Dienekes, not Anthrogenica. Referring to yourself as "us J2s" and listing the things that YOU did, like building the monuments on Malta, is so fallacious, it begged a response.

Mae Kae said...

Can you respond to this please

Hexapolar Autosomal Analysis of the Mediterranean

In Greece it shows it all mixed from Neolithic & later became modern Greeks with less gene flow in historic times, but they say Spain has the most European DNA & Malta has more European DNA than Greece or Italy?

Grogard said...
This comment has been removed by the author.
Kara Moore said...

I just ran the latest Eurasia K11 CHG-NAF Admixture Proportions and this is my results...

Papuan 2.08%
Neolthic_Anatolian_Farmer 22.71%
S_Indian -
Eastern_Hunter_Gatherer 13.69%
Kalash 2.62%
E_Asian -
Amerindian 0.42%
E_African 1.48%
Caucuses_Hunter_Gatherer 17.19%
W_African 0.21%
WHG 39.60%

The known ancestry from that part of the world is Bavaria, my great-great grandparents, and only one set of great-great grandparents. The rest of my family is Colonial American.

22.7% seems to be a huge chunk to me, but perhaps this is the more recent Bavarian ancestry.

So far all the calculators seem to bounce around the whole Alpine and Pannonian Plain region, solidly without fail, Hungarian pops up every time.

My mtDna haplotype is T2b, and that seems reasonable for my mother's ancestry, but her mother and mother and mother and mother were Colonial American. There has been no way to trace back beyond that for me on that line. The Bavarian comes from her father's father's side.

So I have ancestral DNA of Anatolia (before that they were called Hittites) and yet not on my mother's maternal side.

Something must be mixed up, 23 and Me states that I am 46% Irish/Britsh and 23% French/German.

OK,so that's 500 years ago, true enough, I do have ancestry from Scotland and England. Then on the Oracle for the calculator, this is what it says

1 English 3.49
2 Hungarian 3.94
3 Croatian 4.56
4 Scottish 5.1
5 Czech 5.13
6 Ukrainian 7.59
7 Norwegian 7.66
8 Icelandic 8.18
9 Spanish 10.8
10 Bulgarian 12.49

Hungarian??? ...check if Hungarians were Anatolians
Croatian...check only if Bavarians were Croats.

OK, so it is relatively southeastern Europe.

What is the WHG?

But then for 500 years.
Norwiegan..probably Viking
Icelandic...also probably Viking and contributes to English/Scots

Then Bulgarian. That finishes the top ten.

Is WHG, Western Hunter/Gatherers?

I think that new names should be given, Alpine and Pannonian. I have no problem being from either.

But can you please, for the love of God, explain the Papuan? I just don't get that one.

Justin said...

I think the Anatolian and Greek farmers are essentially the same people - stock wise. I don't think there needs a mass migration posited. The culture was likely connected across the Aegean region and perhaps is pre Bosphorus flooding.

mooreisbetter said...

Dienekes is one of the few who has been humble when the (lack of) data requires humility, and confident when the data allows one to make a prediction.

I have posted repeatedly about this concept: how aDNA is solving mysteries and proving or disproving theories wrong, but, how little we still know, and the major gaps. My latest.

I think this concept was one of the few where it was warranted to make a prediction. But there are many others where our knowledge is still very lacking.

Gaspar said...


I have never read about any people stating Hattian and Hurrian adapted and used by the Hittites ~1700BC is anything except indo-european. It is clearly stated as not being from the semitic language tree.
With this knowledge, then the hatti and hurrians are also clearly a non-semitic people mostly likely from the southern Caucasus area.
There are over 50 volumes of hittite script which have been studied in respect to language.

Thomas Alan said...

I think you missed one: ancient mtDNA from Romania confirms early neolithic farmers cluster with LBK but middle neolithic farmers already cluster with modern Europeans. This seems to now be the earliest evidence of a predominantly H haplogroup mtDNA population in Europe, a couple thousand years before corded ware culture, and suggests they were brought in by farmers via Anatolia or perhaps the southwest Ukraine agriculturalist zone, not by pastoralists from the steppe.