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Thursday, October 1, 2009

The Neolithic aDNA map of Europe


Based on: Izagirre & De la Rúa 1999 (Basque Country by the Upper Ebro, huge sample), Di Benedetto 2000 (Catalonia), Chandler 2005 (Portugal), Haak 2005 (Neolithic Central Europe), Sampietro 2007 (Italy), Bramanti 2009 (Neolithic "hunter-gatherers" of mainland Northern Europe) and Malström 2009 (Funnelbeaker and Pitted Ware from Sweden). But I mostly used the Ancient Eurasian DNA site for the references.

For comparison, see my previous post and map, on Paleolithic aDNA. I tried to use the same color coding.

  • Other means N1a in Central Europe, T+X (sic) in the Basque Country and I1 and W1 (one each) in Catalonia.
  • T is T2 where tested downstream, but in some cases this is not determined.
  • U* is U4 in Catalonia and Götland, U3 in SW Germany. Not determined in the other cases. In the Basque Country it should be U5 or U8a attending to modern clades and in Portugal could well be U6 for the same reason.


It is quite hard to deal with, as we have huge blanks, specially in the East and SE. It would seem that Danubian Neolithic (LBK) meant a "Mediterraneization" of Central Europe. The new lineages nevertheless should have been picked up in the Balcans or Middle Danube but we have very limited data for that area. What we do have though (Austria, Hungary, North Italy) is consistent with such idea: 2 H, 1 T, 1 N1a (or 1 H, 1 N1a if you prefer to skip Italy).

It must be said that the importance of the oddball N1a seems somewhat hyped, as most of the cases appear in a small and then rather marginal area of East Germany that just happens to have been oversampled. If we exclude that district, the LBK mtDNA pool has only one N1a case, with T2 (4), H (3) and K (2) being the dominant lineages instead.

A counterpoint is Iberia, where Neolithic data is abundant:

In Portugal, continuity is pretty obvious: H, U5 and U* (U6?) persist at similar apportions as before and the main change is the loss of some "other" lineages and incorporation of novel V ones.

The large Basque sample does not belong to the archetypical mountain Basques but to the populations of the Ebro Valley, where the anthropometric data shows a clear input of "gracile mediterranean" types upon Neolithic arrival. It has nevertheless loads of H (c. 40%), suggestive of a good deal of continuity with the pre-Neolithic populations, if these were similar to the known Portuguese ones. The presence of U in large amounts is also suggestive of continuity and should be U5 and possibly U8a. K, J and that T+X "other" groups (totalling c. 40% too) look of Neolithic arrival, though the place of coalescence of K and T2 has not yet been determined safely.

A similar case seems to be that of Catalonia, though here I miss the mtDNA K, so over-abundant in the Basque case.

Review by lineages:

Haplogroup U:
  • U5 - Clearly Paleolithic, once dominant in Central Europe and maybe other regions of the north but also frequent elsewhere. Today still somewhat common but specially in NE Europe.
  • U4 - Also spotted among Central European hunter-gatherers. Today more or less like U5.
  • K (derived from U8) - First seen in Neolithic Basque Country (lots!) and Central Europe (proportionally also very important), as well as Swedish Pitted Ware (regressed foragers of Chalcolithic Age) and Alpine Italy (Ötzi). Today is widespread at low amounts with highest frequency around the Alps, Bulgaria, Kurdistan and Georgia.
  • Other U - Found in Swabia, Portugal and Morocco in the Paleolithic/Epipaleolithic. In the last two cases it could be U6 (today somewhat common in North Africa and Iberia). U3 first spotted in Neolithic Baden-Würtemberg (today most commonly found in Bulgaria, the Upper Danub and Britain maybe).

Haplogroup R0:
  • HV - Found in Gravettian Italy and maybe other locations. Rare now in Europe but somewhat common in West Asia.
  • H - First spotted without doubts in pre-Neolithic Portugal and Morocco, though it could have been common in Paleolithic southern Europe. Expanded to the North since Neolithic, it seems. Today widespread in Europe but most frequent in the Atlantic coasts and SW, as well as Albania, South Italy, Denmark and Norway.
  • V - First detected in Paleolithic Morocco, in Europe not until Neolithic (Portugal, Central and North Europe). Today most common among Saami (Lapps), sub-Pyrenean areas and parts of North Africa. Also North Italy, Finland and at lower frequencies almost ubiquitous.

Haplogroup JT:
  • JT - First spotted in Paleolithic Morocco. Today JT(xJ,T) is rare and mostly seen in North Africa and Italy.
  • J - First found in Neolithic Basque Country, Catalonia East Germany and Sweden. Today widespread at low frequencies, somewhat higher in Northern Germany. Quite frequent in West Asia. Used to be the archetypal "Neolithic mtDNA marker".
  • T - First detected in Neolithic Central and Northern Europe, as well as North Italy and Catalonia. Rather dominant in Neolithic Rhine area (40%) and Alpine Italy (50%). Seems all T2. Today variedly common, maybe most frequent in North Italy, Georgia and some Eastern European locations.

Haplogroup N1:
  • N1 - Found in Gravettian Italy. Not found as such anymore, I believe.
  • N1a - First spotted in Neolithic Central Europe, being dominant in East Germany (35%) and East Hungary (1/1). Apart of one case in Rhineland, not seen elsewhere. Today very rare.
  • I - First found in Neolithic Catalonia (I1). Today variedly common... or not, at low frequencies anyhow.

Haplogroup N2:
  • W - First detected in Neolithic Catalonia too. Today variedly common, specially in the Mediterranean and NE Europe. More frequent in West Asia and even India (where its cousin N2a is also found) and for me a quite clear "Neolithic marker" too.

Haplogroup X: not detected in aDNA up to Neolithic. The Basque Neolithic sample of Izagirre mentions 3 "T+X" but considering what is found elsewhere in Europe at the time, it's probably all T. Today mostly West Asian (X2), North African (X1) and Native American (X2).


Updated (Oct. 5):

1. Corrected the legend to show haplogroup K.
2. Added Ötzi's K lineage to the series of Alpine Italy, as it's within that timeframe.

Please notice that the dates of the samples vary from c. 5500 to 3000 BCE. The later samples would be considered Chalcolithic by most prehistorians but the Baltic groups with high U were all (regressive?) hunter-gatherers. The dividing line between the various periods is not something straight but I find better anyhow that the maps follow consistent chronological timeframes. In this case, the map covers the period between Neolithic arrival and that of consolidation of Kurgan immigrants (Indoeuropeans), at the early/middle 3rd milennium (Globular Amphora and Corded Ware), event that may well split European Late Prehistory in two.


Anonymous said...

This is from argiedude. Maju, nice maps. You have to add the legend for the blue squares (K, right?) in the Basques, between the green (J) and light blue (U*). I presume the Portuguese Neolithic by Chnadler was part of the same study that tested the Portuguese paleolithic, right? And there's no haplotype data, I presume?

Maju said...

You have to add the legend for the blue squares (K, right?) -

Right. My bad. I also wanted to make both maps the same size and somehow I did not. :/

presume the Portuguese Neolithic by Chnadler was part of the same study that tested the Portuguese paleolithic, right?-

Chandler 2005 reports two sets of data for Portugal: Epipaleolithic ("Mesolithic"), n=9, and Neolithic, n=23 I think. They do not give the haplotype data nor any extended info (it was a presentation for a congress, not a full-fledged paper). I had to determine the number of individual cases by using graph apportions, so minor error (one indvidual misplaced) is not excluded in the larger Neolithic sample (the Epipaleolithic sample is very clear: 5 H, 1 U5, 1 U* and 2 exotic).

They mostly discuss, based on a PC graph, whether the two populations are related or not. They decided they were not but I think it's not fully justified and maybe influenced by Zilhao's position on the extent of change with Neolithic arrival in Portugal on an archaeological reasoning that I may not agree with. They do admit that the source population for Neolithic cannot be West Asian, because it lacks haplogroup J and also that it can't be totally determined wether there is continuity or not, as the Epipaleolithic sample is too small.

My impression instead is that of possible continuity based on that of H and U (including U5). But admittedly I have no explanation, other than chance, for the vanishing of the exotic fraction and arrival of V. Chance cannot be discarded in any case, specially as most Epipaleolithic individuals were from a small district (Sado valley), so it may be a similar case as with the N1a in Neolithic Germany: very localized founder effect/s.

Maju said...

Ok. Fixed the issue of K and also that of different map sizes (don't know why this last happened).