Venneman's exposition of Vasconic substrate in Western Europe

Found at Iruinaoka's page at YouTube, which also includes many videos on Iruña-Veleia, most of them in Spanish language.

However linguist Theo Venneman's exposition is in English, even if his German accent and poor sound quality make sometimes difficult to follow. Turn volume up, adjust balance and be patient... or look for a better quality video/book/paper. I apologize in advance for that poor quality, which is annoying, but I cannot do anything about it.

Video 1: greeting in Basque, introduction on Basque known and speculated history (Upper Paleolithic) and vigesimal system in Basque, Western Romance, Gallic and Germanic:

Video 2: Verbs: difference between to be at/in/on (egon) and to be something (izan). Accents.

Video 3: Loanwords: kinfe (< ganibet), silver (< zilar), star (< izar), andra (lady, woman), grand (< grandi < handi)...

Video 4: More loanwords: cheese (< Käse < gazta < gatz = salt), bust and bush. Toponimy: Lech.

Video 5: Toponimy:  

Is- (Isar, Isère, IJzer, etc.), from Iz- found in compound words like izurde = dolphin ("water boar"), and ur- (from ur = water).  

Aran (valley), from which: Arn- in Germany/Austria, -earn in Scotland.  

Ibaso (ancestor river?) -> Ybbs, Ibar (river bank) -> Eber-, Iberus/Ebro...

Suffix -oz (place?)

Video 6:

Toponimy: -os.

Heint/haint (German): today, from hi naht ('this night'), construction similar to Basque gaur (gau: night + (hau)r: this), also reported by Caesar in Gaulish.

Coherence between linguistics and genetics.

See also:

• Theo Venneman's homepage, 
• B. Oyharçabal's Power Point presentation (including many of the same graphs used by Venneman in his exposition): direct download.
• R. Mailhammer's The Prehistory of European languages (PDF)

_________________________________________

Notes: I must say that I do not necessarily agree with every single etymology but I do with the bulk of them at least. 

I must also say that I miss mention of some Vascoid roots I considered on my own, some very apparent, for instance:

• The obvious bi- Latin particle for two or double (as in bilateral, bipartite, etc.), which is not Indoeuropean and is terribly consistent with Basque bi: two.
• Professional suffix -er/-ero, very similar to Basque -ari, which is used the same way but has a clear Basque etymology: ari (auxiliary verb of action, used in present/past continuous), arin (fast, quick), aritu (to hurry).
• English verbal infinitive particle to (as in to be, to do...), which sounds identical to the most common verbal infinitive suffix in Basque: -tu (sometimes found as -du for cacophony avoidance). Other verbal endings are -n and -i. This one is a bit conjectural. 
• English ash, Basque auts (ash, dust).
• English kill and ill, strikingly similar to Basque il(-du) (pronounced like ill), meaning 'to die' (intransitive) or 'to kill' (transitive).

 More conjectural:

• English black, compare with Basque bel-(tz) (black), which is at least coincident in the two main consonants. However there is a Germanic-IE etymology as well from *blegh (to burn, shine) but this is the same root as blanco/blanche (white) and blank. Conjectural.
• Arguros and argentum, silver in Greek and Latin respectively. Possibly from Basque argi (light, shiny/bright, to shine), would be a loanword from the Bronze Age possibly. The only Eastern IE word is Sanskrit arjúna (not silver but white) but may well be unrelated or a borrowing via Mycenaean Greek. Conjectural.
• Possibly mountain (V. Lat. montanea) and mound. While this is argued to be an IE word, Eastern alleged cognates appear without the intermediate -n- and mean different things (such as shore in Albanian mat). I conjecture it may be from Basque mendi (mountain) instead, in turn from mende (power, might), rel. mendebalde (West: 'part under power' or maybe 'under the mountain') and possibly from a very archaic West Eurasian root *man, as in manna, man, Lat. manus (hand), meaning power or potency.
• Coincidences of declined to be: Basque zara (you are), English are, Spanish eres (you are). Also notice the similitude between IE *is (to be) and Basque izan (to be). Also coincidence in the second person but dancing between singular and plural: Basque zu (singular 'you' but distinct from hi: thou), English thou, German du, Latin tu. These and maybe others might point to a common very old origin of Basque and IE or to the affection of PIE by a Basque-related language such as NE Caucasic/Hurro-Urartean/Sumerian (conjecturally the language of Eastern Gravettian). Alternatively some may be influences of IE into Basque but most look as deep phylogenetically rooted in Basque.

R1b1b2a1 is almost unique of West Europe

[Typo: in the maps M529, also known as L21, is wrongly written as M259. My apologies]

This is one of the virtues of Myres' paper (that I mentioned yesterday): that a somewhat more clear phylogenetic subdivision is made, emphasizing the difference between West European R1b1b2a and other R1b or R1b1b2, often blurred in previous papers, causing great confusion even to researchers themselves.

A defect is that instead of using a standard name for the defining SNP (L51/S167 per ISOGG) they chose to name it M412. However Argiedude says it is the same marker and I imagine it is. [Update: confirmed: the rs number indicates it is the same SNP].

Another virtue is that some of the substructure of R1b1b2a1 is also mapped, what really covers pretty well the Northern and Italian area of spread of this lineage and even some relatively unmapped areas of SW Europe, specially France.

In any case the apparent structure is curious, so I got the supplementary table S4 (supp. material is freely accessible) and made this map:

Click to enlarge

Notice that, following Argiedude, M412 stands for L51 and M529 stands for L21, what, if confirmed would make the following equivalences:

• R1b1b2a-M412 = R1b1b2a1 (L51)
• R1b1b2a1a2-M529 = R1b1b2a1a2f (L21)

Even if not confirmed, the equivalence should be approximate anyhow.

I'm sorry for the horrible color palette but it's my first attempt to make pie charts with Open Office spreadsheet gadget. Next time I'll do better I hope.

Notice also that I did not use all the samples, in the cases of small countries or less relevant regions I arbitrarily chose and discarded some.

Finally notice that pie charts represent only apportions of R1b1b2 and say nothing of the frequency of the lineage overall, which in most of East Europe and West Asia (excepting Turkey and a few neighbors) is extremely low.

The apparent structure of R1b1b2a1

The most apparent structure is, as we already knew the rather different R1b1b2a1a1 and R1b1b2a1a2 distribution. The first one (color coded brown and light blue) is dominant in the North and rather rare in the South - hence: 'North clade' for short hereafter. The second one (dark green, light green, purple and light orange) is by comparison not just more frequent in the South but also probably more diverse as well - hence: 'South clade'. However it is also found in the North.

Then there are some transitional "remnants": R1b1b2a1a* (L11) and R1b1b2a1* (M412). These should be informative (meaning some extra diversity at their structural levels) in order to infer the history of the haplogroup.

Per the hierarchical distribution seen here and diversity data from older works, the most likely origin of R1b1b2 as a whole is Anatolia.

Then R1b1b2a1* (M412) (yellow) is suggestive of a Mid-Danubian (or Italian or Iberian) coalescence.

R1b1b2a1a* (L11) (middle green) is suggestive of a West-Central European (or SW European) coalescence. More data on the Pyrenean region would clarify this maybe.

And, after this layer, comes the division into the widespread Southern and Northern clades mentioned before.

A reasonable interpretation is that the lineage traveled relatively fast upstream of the Danube (and/or via North Italy onto the SW), branching out then into the two major North/South clades. These two lower level lineages are in fact the two main stars of this demographic expansion.

My bet is that this represents a wave of colonization of Europe (when?) with secondary expansions from SW Europe (Franco-Cantabrian region possibly) and Central Europe (Rhine-Danube region I presume). There are several scenarios that can account for this, essentially Paleolithic (or pushing into the Epipaleolithic).

I don't see clearly how this structure could account for a Neolithic spread, really: no Mediterranean Neolithic pattern is apparent at all and Danubian limited expansion cannot account for any spread to SW Europe, certainly not South of the Loire and certainly not at the frequencies it is found there (nor in Britain/Ireland either). Claiming a Neolithic spread of R1b1b2 with this structure can only be done from a very shallow understanding of Neolithic archaeology and prehistory overall.

The main known demographic expansion we know of in the European Upper Paleolithic is the one after the Last Glacial Maximum, when Magdalenian culture expanded from the Franco-Cantabrian region, both northwards to Central Europe and, later, southwards into Iberia. This led to cultural divergence into the Epipaleolithic, with the important expansion into the newly available areas of the Far North, earlier covered in ice. Within the Epipaleolithic some further cultural flows are detected: from the Franco-Cantabrian region into Iberia (Azilian) and from somewhere in Mid-West Europe into the Southwest (Sauveterre-Tardenoisian).

Also, back to the LGM, Magdalenian techno-culture may have got a NW European ultimate inspirational origin but anyhow mediated by the warmer and richer Franco-Cantabrian region, where the culture flourished properly.

It's difficult to reconstruct in detail but, as far as I can tell, the two main North/South clades must have expanded in the Magdalenian period (one from the Franco-Cantabrian region, the other from Central-NW Europe itself) and also in the ulterior Epipaleolithic. Neolithic does not seem able to account for much but may have helped to shake the board a bit, specially in East-to-West direction.


Frequency maps

Selected frequency maps from the paper
Click to enlarge

Notice that the expansion of the South clade to the Atlantic islands does not invalidate its southern character and probably represents an Epipaleolithic-to-Neolithic spread.

Notice also the large amount of unclassified Southern clade in Iberia. The area around the Pyrenees was not really sampled in this study and therefore it is distorted by neighbors ("South France", looking more like SE France, Valencia and Cantabria).

In order to appreciate better the real thing in this aspect it's probably good to take a look at Cruciani 2010, who did bother to sample near the Pyrenees and gets maybe better (or at least complementary) maps illustrating the same problem.


Update: I superimposed (with complementary colors) the South (red) and North (blue) clades from the frequency maps above. However in order to account for the differences of frequency, I had to lighten the blue shade (North clade) because the scales are different. Take it as an "artist's impression" anyhow:



Update (Aug 27):

Here there is a hopefully better version of the map at the beginning of this post:

Click to enlarge

I put special care in giving each distinct clade an specific color range for easier visualization. All R1b1b2a1 (M412/L51/S167) seems to have coalesced in the Central-to-Western European area but the real expansion seems to have happened after this haplogroup split in two, which I dubbed the North and South clades.

And this is my reconstruction of the haplogroup expansion:

Click to enlarge
Color coded as above

Update (Aug 28):

Take a peek at the comment section, where I briefly discuss molecular clock difficulties and also the only possible Neolithic scenario for R1b1b2a1a2 (South clade): a massive demographic expansion in the context of Megalithism.

Rejecting or confirming this would require greater research in the structure hidden "under the asterisk" in SW Europe. At the moment only two minimally-sized sub-haplogroups are known: Basque/Gascon-specific R1b1b2a1a2b and sub-Pyrenean R1b1b2a1a2c (Gascon, Catalan, etc.). This alone gives highest structural diversity to the Pyrenean region, however most of the South clade remains unresolved (hidden under the asterisk), both in the Pyrenean area as in Iberia proper. And the key issue to solve would be if R1b1b2a1a2 is most diverse at the Pyrenees, what favors a Paleolithic spread scenario, or in West Iberia (and Brittany/West France), what would favor a Neolithic-Megalithic spread scenario instead.

Also it's maybe important to remind here the excellent STR work of Laura Morelli earlier this year, which was discussed in this article.

Importantly, this graph (annotated by me):


The graph is suggestive of the existence of another "West Asian" distinct haplogroup "under the asterisk" (that I labeled "R1b1b2a2?") and a possible Balcanic, rather than Anatolian origin for the R1b1b2 clade.

If so, this would correlate with the high diversity of the (much smaller) brother haplogroup R1b1a in the Italy-West Asia arch (as well as in Central Africa) and would suggest a slightly different origin and scatter for R1b as a whole (ref 1, ref 2).

Megalithic aDNA from Charente Maritime

The paper requires payment and the abstract is not too informative but John Hawks mentions in his blog the essential findings and Jean Manco has already added its results to her exhaustive aDNA list.

Testing only for HVS-1 (hyper-variable control region 1) haplotypes, they found that three individuals buried in Megalithic context at Prissé-la-Charrière, near La Rochelle (Charente Maritime, France), dating to c. 4200 BCE, have the following mtDNA haplogroups: X2, U5a and N1a.

Notice that HVS-1 sequencing is subject to potential criticism because, sometimes, different haplogroups may share the same or very similar signature. Generally aDNA studies are considered more reliable when they go to great lengths to make sure that there is no contamination or sequencing error (they seem to have accounted for that in this case) and when they can confirm haplogroup adscription by testing for SNPs in the control region of the mitochondrial genome.

In any case, this is more data to add to the list of European aDNA, on which I may work out an actualized synthesis soon. It is also intriguing data.


N1a

The authors, as Hawk and others, focus specially on the presence of N1a and with good reason: this haplogroup, now residual among Europeans, has been showing up, sometimes at high apportions, in aDNA from Central European Neolithic sites belonging to the Linear Pottery culture (Danubian Neolithic, LBK by its German acronym): 1/1 in Hungary, 4/11 in East Germany and 1/10 in West Rhenania. The haplogroup nowadays is most common in West Asia and had not been found before in any European aDNA other than Danubian Neolithic and some later Hungarian ones.

It is also noticeable because its relative haplogroup I (also part of haplogroup N1) has been shown to be relatively common among ancient and medieval Danes but is now quite rare.

While other haplogroups like K or various subclades of U also seem to have experienced some contraction, their decline is nothing compared with that of N1 sublineages.


X2

Haplogroup X2 is also somewhat noticeable in my opinion. The lineage is not that common either in Europe (excepting to some extent Orkney islands and some Mediterranean scatter) and can easily be speculated of as a Neolithic arrival. However the antiquity and wide spread of this lineage (from Europe to Native Americans via Altai) does not allow us to reach to conclusions easily.

In any case this is the oldest individual known to have carried this haplogroup. Excepting some unclear cases from the Southern Basque Neolithic (T or X), the next case are two siblings from a Corded Ware culture burial in Eulau, East Germany, c. 2600 BCE, and then some scattered findings from historical times.

This haplogroup, unlike N1 (N1a and I), does not pose any real issue because it is not really being found at high frequencies at all, just like today.


U5a

There seems to be some confusion about which haplogroup is this one: while Hawks reports it as U5a, Jean Manco does as U5b. However searching for the sequence provided by Jean herself at PhyloTree, I can only find a close match in U5a, probably a variant of U5a2a1.

This is where the sequence 16270T, 16271C leads me to. However there is another marker not defining this U5a2a1 haplogroup, 16189C, but this seems extremely variable, with matches in every other haplogroup in the mtDNA tree of humankind, so it is very possible that it only responds to individual or microlineage variance in this case.

U variants including U5 (and U2, U4, U*, etc.) have been reported in the most ancient DNA sequenced anywhere in Europe: Kostenki culture in Russia (1/1 U2), Gravetto-Solutrean in Andalusia (1/2 U*), Magdalenian in Swabia (2/2 U*) and the Epipaleolithic of Portugal (1/9 U5, 1/9 U*), the Don Basin (2/2 U5a), Swabia (2/4 U5b2, 1/4 U5a, 1/4 U4), England (1/1 U5) and Lithuania (3/4 U5b2, 1/4 U4). Additionally they have been reported as dominant in subneolithic or regressive neolithic peoples in the island of Götland and related cultures in Baltic Poland and Germany (Pitted Ware senso lato).

Later the presence of haplogroup U and specifically U5 becomes less common, although it keeps showing up until present, being nowadays a most important haplogroup in all Europe, specially towards the NE. In that part of Europe we do find not just U5a but also U5b and U4 at relatively high frequencies. This is largely coincident with what we do find in aDNA, with the caveat that everywhere but Iberia, the haplogroups seem to have shrunk somewhat since Neolithic arrival (what is easily explained in terms of partial population replacement).

However there is much more than that to haplogroup U: U2 for instance has a more Asian distribution, U6 is typically North African and Iberian, being extremely rare elsewhere, U3 is typical of the Black Sea area and the most common subclade of U8, K, is very much widespread nowadays but only shows up in aDNA since the Neolithic (Alps, Central Europe, Pyrenees). Haplogroup U after all is a very old macro-lineage, by all accounts dispersed at the very colonization of West Eurasia some 40-50,000 years ago.


In context

Neolithic mtDNA in West Eurasia (updated)
Each dot represents one individual
High U Baltic area sites are subneolithic or regressive Neolithic (Pitted Ware)

Hawks, following Dienekes, argues that:

If N1a were present somewhere in pre-Neolithic Europe, it would require some kind of "partition" of the pre-Neolithic population, along with its propagation -- presumably southeastward -- into the LBK of central Europe. Seems doubtful.

I fail to see the point, sincerely. It does not require anything of that, I don't see why. We must remember that Neolithic diffusion, following the archaeological record, is a complex process in many cases showing clear signs of cultural (and hence demographic) continuity at least in the Mediterranean (Cardium pottery and related remains are often found along locally rooted Epipaleolithic tools and only in specific sites colonization seems the most likely explanation for its expansion). Even if population replacement may be a more likely case for Central Europe, there is still the unsolved problem of where did the core immigrant population originated: West Asia, Greece, Hungary or a mix of all three? The lack of aDNA for the Balcans and Turkey does not really help, the only reference remaining being the PPNB site of Tell Halula, at the Upper Euphrates.

This site does show abundance of mtDNA K and T (specifically T2b) and some presence of the ubiquitous H too, together with L2, R* and C1. The K+T combo shows up also in Central European Neolithic but the explanation for the origin of other associated haplogroups (J and N1a specially) is not clear on light of the present data. It can be argued for them having a West Asian origin certainly but the case is far from settled. The other LBK haplogroups, found in Paleolithic individuals from Europe and Morocco, can hardly be argued to have spread with Neolithic (the problem of the expansion of H to Northern Europe remains murky however with my favorite explanatory vectors being either the late Bronze Age Urnfield culture or Megalithism but notice also low Paleolithic/Neolithic sampling in the North Sea area, still allowing for UP presence for this lineage). The causes of the apparent decline of N1 (N1a and I) in Europe remain equally confusing as well.

The particular case of Prissé-la-Charrière also requires an explanation but I can only give a negative one: it cannot be attributed to LBK in any case because Danubian Neolithic never reached so far south. But it actually contrasts more with the samples related to Cardium Pottery (Mediterranean Neolithic) and Megalithism (largely "Atlantic Neolithic") in Iberia and Italy, which display a very different array of haplogroups (H, U*, U5, K, J, etc.) This in turn makes more difficult to argue for a Mediterranean or Atlantic Neolithic origin of this localized aDNA sample.

The case remains open in my opinion.


References:

Marie-France Deguilloux et al., News from the West: Ancient DNA from a French Megalithic burial chamber. American Journal of Physical Anthropology, 2010. Pay per view.

Building History: Ancient Eurasian DNA.

Leherensuge: European ancient DNA in sequential maps (not updated in almost a year).


Update: Link to the whole paper (PDF), courtesy of Natsuya. They report the U5 individual as U5b, by the way.

Archaeology versus some bad Genetics

I just stumbled upon a somewhat old but very interesting article on why population genetics has gone mad and leaned towards Neolithic replacement against all archaeological evidence. Why? Because they never cared at all about the archaeological evidence, not even about the genetic evidence that contradicted their own conclusions: they have been acting ideological all this time. Sadly enough.

Mark Pluciennik, Clash of Cultures? Archaeology and Genetics. Documenta Praehistorica 2006. Freely available PDF.

Worth a read, really.

Finally some good research on R1b1b2!

This is a most important paper for those interested in Y-DNA R1b and specifically European and West Asian R1b1b2:

Laura Morelli et al., A Comparison of Y-Chromosome Variation in Sardinia and Anatolia Is More Consistent with Cultural Rather than Demic Diffusion of Agriculture. PLoS ONE 2010. Open access.

An excellent critique of Balaresque 2009, totally in the line I was favoring myself.

Most important is maybe fig. 2A, where the use of (novel?) STR marker DYSA7.2, in addition to the usual ones, unveils a clear structure within R1b1b2-M269. I have annotated it for easier visualization:

Click to enlarge

Duality

The already quite obvious Europe-Anatolia (A-B) duality of this haplogroup becomes terribly apparent: two different star-like structures connected by a less clear group (C) centered at the Balcans.

I imagine that sooner than later, someone will come up with a defining SNP for the more than likely Anatolian haplogroup. If I'm correct, with the current nomenclature it should be something like R1b1b2a2. However the root of the whole R1b1b2 haplogroup might also be in that cluster (or is it in C?).

It is surely important to notice that this presumptive R1b1b2a2 haplogroup (at least a clear haplotype star-like structure) also experienced quite apparently a rapid expansion of its own, mostly in West Asia and the Balcans, which is in turn surely related to the expansion of R1b as such through the Mediterranean, Africa (R1b1a) and Central Asia (R1b1b1).

R1b expansion pattern

We may well say that R1b expanded quite dynamically but not with total simple continuity but in colonization "bouts". The first bout is probably best represented by R1b1a scatter in the NE Mediterranean arch (two subclades) and around Central Africa (maybe with a Sudan origin, two subclades too). The second bout is surely a minor one towards Central Asia represented by R1b1b1 (typical of Uyghurs, an odd Chalcolithic founder effect in a previously desert area). The third bout is the spread of R1b1b2-M269 in West Asia and Europe, which represents the bulk of the R1b worldwide in raw numbers.

This third bout is actually two, as is evident in this paper (but was also evident before for those who can "read between lines" at least) but they may have happened in parallel (in which case my annotation above is more strictly correct) or they may have happened sequentially (in which case, lineage B would be R1b1b2a*, except for one of the branches that is actually the root: R1b1b2*).

A quite plausible reconstruction of the spread of R1b (c. 50-30 Ka ago?)
Click to expand

Synthesis

Whatever the case, two things are clear:

1. There is a neat distinction between R1b1b2(xR1b1b2a1) and R1b1b2a1.

2. R1b1b2a1 had its own distinct and fundamentally unique expansion in West Europe, sensu lato.

Also notice that, as happens in previous versions, there are many branches hanging from the R1b1b2a1 central node (modal haplotype also), which appears to contradict the known subhaplogroup structure that only recognizes two branches and which seems to be quite imperfect as of now.


Tempo

As you know I am not any fan of the molecular clock but it's worth mentioning the estimates of this paper for they totally contradict the Neolithic model: the pool of Sardinian and Anatolian R1b1b2 gets an estimate age of 32.6 Ka, the Sardinian one has an age of 27.0 Ka and the Anatolian of 19.6. But the regional ages may be misleading because they include members of both clades, however the overall age should be roughly correct within the molecular clock paradigm.

This would fit with a Gravettian expansion but could also fit with an even older Aurignacian one. It makes some good sense, specially as we are talking of a single expansion that replaced all what was there before, which could only happen at very low population densities, and also because it is a single expansion and not two. It also makes good sense with the likely pattern of expansion of its likely mtDNA companion haplogroup H (also V probably).

As the authors admit, the Magdalenian late glacial recolonization (and noticeable demic expansion) is also within the scope. However I find difficult to explain R1b1b2a1 in Italy within this context, as Italy remained Epigravettian until the very Neolithic. It is also much easier to replace all when there's almost nothing to replace, as was surely the case after the Campanian Ignimbrite supervolcano eruption of c. 40 Ka ago.


Other lineages

The paper is also of interest for other haplogroups such as E, G and J. However I fail to see anything clear surely because of the excessive emphasis in exploring Sardinian-Anatolian connections without a wider scope.


See also: for a completely opposite editorial opinion and likely heated discussion at Dienekes'.

Again on Neolithic and European Y-DNA

Specially on why R1b1b2a1 (the bulk of Y-DNA R1b and the Western European specific subclade) cannot be Neolithic. It comes up in all discussions and I really feel the need to explain: to create a generic post that serves as reference.

For that purpose I created a simplified map of European Neolithic cultural flows (excluding mostly Eastern Europe that anyhow had its own distinct processes):

Click to expand


The Early Neolithic

As you probably know, European Neolithic (defined by the existence of farming and animal husbandry, and also often of pottery) began by all accounts in Thessaly, Greece (Sesklo Culture). It's characterized by a pottery painted using mostly the colors red and white and with those colors it spread through the Balcans.

More intriguing are the origins of the Cardium-Imprinted Pottery culture, specific of the Western Balcans, who did not use color in their pottery and instead decorated them with patterned impressions often made with the shell of a mollusk of the genus Cardium (hence the name). Some early pottery of this kind has been found at the Thessalian site of Otzaki, along with finer pottery of Sesklo style, however it is only known as distinct culture once it became dominant in the Western Balcans.

This Balcanic duality defines the general duality of early European Neolithic because Cardium Pottery Culture would spread by the Mediterranean shores, largely by boat, while the main Balcanic Neolithic would eventually give rise to the Central European one, often known as Danubian Neolithic.

The Cardium Pottery Culture spread with some colonization but mostly by acculturation of the indigenous peoples, as is evidenced in the toolkit continuity in most sites. They were specialized in fishing and sheep/goat herding but also carried the full package of cereals and at least one legume: lentils. They built villages but also inhabited in caves. After the initial coastal expansion local expansions took place as well already under the tag of Epicardial, most importantly along the Rhone and Ebro rivers.

Meanwhile in NE Hungary (curiously the easternmost reaches of Epi-Magdalenian culture) an ill explained cultural shift happened: the Balcanic pottery style was replaced by an engraved one. It is the Eastern Lineal Pottery Culture, which had a very limited expansion on its own (mostly to Transylvania).

However a derived culture arose then in Western Hungary, Moravia and Eastern Austria: it is the Western Lineal Pottery Culture, more commonly known as Danubian Neolithic. This one had a massive spread through Central Europe reaching into Belgium and Northern France (where it shared the territory with other minor local Neolithic cultures for some time) by the West and into Moldavia by the East, where a previous local Neolithic culture was assimilated as well. It even influenced the area of Vallachia-Bulgaria at a later time, generating which was maybe the first large European state: the Karanovo VI-Gumelnita culture, older than dynastic Egypt but short-lived.

Now it seems also clear that late Danubian peoples from Northern France brought agriculture to Britain, simultaneously to Megalithic peoples from Western France.

But I'm going too fast.


The Late Neolithic

A key an often ignored region in the second Neolithic phase is SW Iberia (mostly southern Portugal). Here there was also an important cultural shift: the custom of clannic (collective) burial in megalithic tombs known as dolmens or trilithons appeared just a few centuries after the arrival of the first Neolithic influences. Dolmenic Megalithism in this area is older than anywhere else by at least a thousand years.

Only much later, c. 3800 BCE, this custom appears in Brittany and other areas of Western France. In the following centuries it would spread through all Atlantic Europe and many parts of the Western Mediterranean, as well as penetrating in parts of the late Danubian cultural area.

It has been speculated that cod fishing may have been related to this expansion, however there must be more than just a mere economic activity: Megalithism had clearly a cultural, probably religious, element to it.

Many areas of Atlantic Europe only reached Neolithic with the arrival of Megalithism or a few centuries before it.

Another key region is more obscure: Denmark and neighboring areas. Here there was an important seagoing culture known as Ertebölle, which has been claimed to be Neolithic... only to see others rejecting that claim. This is a key issue that should be clarified in order to understand the Nordic Neolithic.

Related to this issue is that of Funnelbeaker Culture (often named by its German acronym of TRBK), which is clearly Neolithic in the North and West, where it is also Megalithic but is not in the East (Mecklenburg, Northern Poland). Some had theorized that Funnelbeaker began in Denmark, derived from Ertebölle after the arrival of Eastern influences related to Pitted Ware (another allegedly hunter-gatherer culture of Neolithic times, seemingly rooted in Eastern European Neolithic), but this is contested by others who think it's derived from some northern offshoots of Danubian Neolithic.

The issue is very murky so I prefer to leave it as it is: a mere anotation of unsolved complexity.

All this sums up pretty well the essence of Neolithic in Europe (excluding the East), I believe. In order to synthesize I had to ignore many local groups, often interesting but that add little to the global picture.


Can R1b1b2a1 be Neolithic?

Many people seem to believe that, in spite of this haplogroup being most concentrated in the westernmost reaches of Europe and decaying towards the East.

The most biased ones love to oversimplify and ignore R1b1b2a1 altogether, talking instead of R1b1b2. This was the case with the recent paper by Balaresque et al. which produced the following haplotype structure, which I have duly annotated to indicate what is not R1b1b2a1 and that way unveil the truth:


Other means West-Central-North Europe
Click to expand

It is obvious that the star-like expansion of R1b1b2 happened already at the level of R1b1b2a1 and happened already in Europe: either in Central or West Europe.

It is obvious that it does not show two distinct centers of expansion nor two different waves as should be the case would it have any relation with European Neolithic spread but instead shows one and only one general expansion affecting essentially to West and Central Europe without any kind of pattern, at least not one that we can easily spot (thank Balaresque for that lousiness).

It is obvious that the center of expansion is not in the Balcans either.

The R1b1b2a1 star-like structure has 15 basal branches, of which at least half show a clear strong presence in Iberia (sample that does not include Basques, as the only Basque sample used by Balaresque was from the North and hence included along all French in the Other category). This alone would suggest that the origin of the expansion should not have been too far from Iberia.

Oddly enough, this graph clearly supports the hypothesis of R1b1b2a1 having expanded with Magdalenian culture from the Franco-Cantabrian refuge in the late Upper Paleolithic.

However I do not bet all my money to this hypothesis because the limited haplotype diversity data I manage rather suggests slightly higher diversity in Central and even Northern Europe (this last only with small samples).

But I do bet all my money to the haplogroup having expanded before Neolithic. There is simply no way that the rapid expansion that star-like structure so clearly indicates could have been caused by Neolithic cultural flows. These can explain the scatter of other lineages like E1b1b1, J2b, G2a, T and even some subclades of I maybe. But that's about all.


[Note: what follows is an update, some 10 or 12 hours after the first publication]

What about Megalithism?

The only possibility for R1b1b2a1 to have expanded within Neolithic would be within the frame of Dolmenic Megalithism. This would fit reasonably well with the haplogroup's spread area, with some notable exceptions and would allow for Portugal to act as a transition zone between R1b1b2a* (that has high diversity in this country, comparable to that of Turkey or Italy) and R1b1b2a1a.

However this poses several problems:

First, while Megalithism may be associated to the origins of agriculture in some areas, it is certainly not the case in most. Second, Megalithism shows high cultural diversity and appears related to many different local cultures: it is not a monolithic phenomenon at all, what should be the case if it was essentially one of demic expansion. Third, some crucial areas do not fit well: East and Central Iberia were never Megalithic but are high in R1b1b2a1, instead North Africa was and shows very low levels of the haplogroup (these are just two examples: Central Germany, Austria and Italy also contradict the pattern). Fourth, Portugal has rather low levels of diversity for R1b1b2a1.


The mtDNA control

As mentioned, R1b1b2a1 has a marked star-like structure, meaning rapid expansion from a single center. Several European mtDNA lineages also have such structures. The most notable one is H, which is the second largest star-like structure in all the human mtDNA tree, after M, having 34 basal sublineages. Its descendant H1 also has a noticeable star-like structure with 15 basal sublineages. Less impressive but still meaningful are their ancestor HV (6 branches), their cousin V (9 branches), H3 (7 branches), H1b'f'g'k'q (5 branches), H2a (5 branches). In the U haplogroup there are also several with presence in Europe: K1a1 (9), K2a (6) and U5b3 (6). Other haplogroups important in Europe with some star-like structure are: T2 (7) and its descendant T2b (6), as well as I (5).

But only one group of those mtDNA star-like structures seems to be parallel in geography and dimensions to that of R1b1b2a1: H and its descendants.

In my opinion, H must have spread in Europe early on, probably with the colonization of the continent by our species in the early Upper Paleolithic. Only that can explain the the massive star-like structure, the high diversity in Central Europe and the known presence of this lineage in Paleolithic Portuguese and Moroccans. I also get those time frames using a control region mutation count. However there is one significant difference with R1b1b2a1: H is also found in Eastern Europe and Central Asia in large amounts, while R1b1b2a1 or R1b1b2 is not (and only R1b1b1 is in in Central Asia). This situation also happens in North Africa, where SW Europe-derived mtDNA H and V is very common (c. 25%) while Y-DNA R1b is very rare.

One possible explanation could be that there has been Y-DNA sweeps in those areas which did not affect so much to mtDNA. This could be because of Capsian Culture in North Africa (Epipaleolithic and Neolithic - Afroasiatic languages) and because of Kurgan migrations in Eastern Europe (Chalcolithic - Indoeuropean languages).

However it is still a weak point, admittedly.

So are there other options? I have flirted with the expansion of mtDNA K, which is a much recent expansion than that of H. But K only amounts to 6% of all Europeans, while R1b1b2a1 makes up more than 50% in all Western Europe. They are simply not comparable. Same for the other potential candidates mentioned above.

So at the moment I think that the expansion of R1b1b2a1 must have happened within that of mtDNA H, what implies a Paleolithic time frame.


Isn't a Paleolithic time frame too old for R1b1b2a1?

As you probably know, I strongly distrust molecular clock age estimates, favoring instead an holistic logic, inclusive of all genetic and archaeological data available.

The earliest Eurasian ancestors of R1b (F, IJK, K and MNOPS) must have been there in the time of the main Eurasian expansion. F, IJK and K surely coalesced in South Asia, while MNOPS did in SE Asia. Y-DNA P represents the only known back-migration to South Asia of this macro-haplogroup.

Y-DNA R and probably R1 as well surely coalesced in South Asia then but R1b already did in West Eurasia (either in West Asia or Italy). This R1->R1b migration must have happened within the colonization of West Eurasia by H. sapiens, which happened in the 50-40 Ka time frame by all accounts. So I presume that R1b is 50-40 Ka old.

Then R1b split into R1b1a and R1b1b, which must have existed in the Eastern Mediterranean arch (Italy-West Asia).

Then R1b1b split into a minor Central Asian haplogroup (R1b1b1) and the main West Eurasian one (R1b1b2). This one shows a clear origin in Anatolia-SW Caucasus, where it is most diverse, specially once excluded R1b1b2a1 (however notice that Italy and Portugal have similar diversity levels, what again makes me wonder about the exact role of Italy in particular on the spread of this lineage), and where the main R1b1b2a node seems to have coalesced on light of its non-R1b1b2a1 scatter.

And then R1b1b2a1 spread in a very quick expansion, not from Anatolia but from somewhere in West or Central Europe. This might have happened either in the early colonization of Europe (Aurignacian culture, c. 40 Ka ago), in the second wave of Gravettian culture (Cro-Magnon type, c. 30 Ka ago) or in the post-LGM recolonization from the Franco-Cantabrian refuge (Magdalenian culture, c. 15 Ka ago). It is really difficult to determine which wave but, if R1b1b2a1 has to correspond with mtDNA H, then the latter has to be discarded.

And that's what I can say on this matter. Really, without a time machine or accurate aDNA testing, we cannot say much more.

The Neanderthal-Sapiens transition at Cova Gran (Catalonia)

Found at Pileta de Prehistoria.

J.M. Moreno et al., The Middle-to-Upper Palaeolithic transition in Cova Gran (Catalunya, Spain) and the extinction of Neanderthals in the Iberian Peninsula. Science Direct, 2010. Pay per view.

A good article can be found at El País in both Spanish and English.

Most relevant seems to be that the transition between the occupations by the two species is separated by a thick sterile layer, meaning that there was no sudden replacement but maybe just the colonization of an empty land.

Cova Gran is located at the Catalan Pyrenees in the province of Lleida.

R1b1b2 and R1b1b2a1 distinct STR diversity

I'm borrowing here the work of Aargiedude (again) because it's such a priceless addition that it really deserves to be paid some attention. If professional geneticists would be half as serious as this amateur, we'd know a lot more and a lot better about our the population history of humankind by now.

These two maps represent the haplotype diversity of R1b1b2a1 (ht15) and R1b1b2* (ht35) respectively:

Click to expand

Aargiedude's own observations at Dienekes' blog follow:

I've finished estimating ht15 and ht35 diversity, it's taken me 2 days to do this, and I think the results are amazing. The diversity clines of ht15 and ht35 are almost polar opposites. I think these results seriously call into question the conclusion of the Balaresque study, which is what prompted me to look into this. There's no surfing-on-an-expanding-wave phenomenon occuring with ht15. It has its lowest variance in the supposed origination point: Anatolia.

Instead, as Maju pointed out brilliantly, the study's tree diagram of their R1b1b2 haplotypes seems the result of 2 separate events, not a single wave diffusion. And he was absolutely right.

The diversity of ht35 doesn't form a gradually decreasing cline. It seems to be uniformly similar from Iran to west Iberia, or at least up to Italy, because there are issues with the validity of the North African and Iberian data (small sample size in one case and confusion with ht15 samples, in the other). Its cline seems to be more north-south than diagonally from southwest to northeast. East European countries have the same ht35 diversity as West Europe, with the special consideration of the west Iberian results.

Some technical details to keep in mind. Ht15 can be differentiated from ht35 by barely 2 markers: 393 and 461. Few studies test 461, so most of the samples I used were chosen on the basis of 393 alone. But about 3% of ht15 and 10% of ht35 have the "wrong" value, becoming confused with the other group. This usually doesn't matter, exceot in countries where there is an overwhelming ratio difference between the frequencies of both groups, such as in Iberia, France, Britain, Netherlands, Anatolia, and the Levant. In these extreme cases, I've included, where possible, 2 pair of results. The top pair uses samples predicted as narrowly as possible, by using both 393 and 461. The bottom pair is the standard prediction using just 393. The top pair should be more accurate, but they tend to lack in sample size, so then again, maybe not. Notice in the case of Iberia, that the less restrictive result changes drastically from the more restrictive result, and results in identical values to Iberia's ht15 diversity estimate, suggesting most of the samples are in fact ht15 samples that are being confused for ht35 because they had a mutation in 393 to the modal value of ht35 on that marker. Curiously, this didn't happen in France, where I was only able to use the less accurate method (393 alone), and yet the result is notably low and different from France's ht15 diversity. I'd seriously take North Africa's high ht35 result (0,30) with a military-issue teaspoon of salt, it's just 5 samples. On the other hand, it's notoriously high ht15 diversity (0,28) is pretty solid.

To recap, Baralesque and all geneticists are stuck in a time warp, they're back in 2003, thinking R1b is just R1b. What a waste, after going through all the effort of collecting and processing the samples, to not have had the sense to test for a few extra key mutations that define some major subdivisions of R1b1b2 and are well known for more than 5 years. The conclusions they reached would then have been very different.

The Balaresque paper was discussed at Leherensuge a few days ago.

R1b1b2a1: Neolithic or what?

I found at Dienekes the reference of a new paper on R1b1b2, focused on demonstrating (quite forcibly) that the lineage is Neolithic and not Paleolithic. Dienekes is, of course, happy that they chose to use his favorite (but rather disliked in the field) molecular clock methodology: the one based on the pedigree mutation rate, which make all haplogroups look extremely recent.

Patricia Balaresque et al. A Predominantly Neolithic Origin for European Paternal Haplogroups. PLoS Biology 2010. Open access. I'm provding a link to PubMed Central because at the time of writing this the original link remained broken.

I am not really happy: the pedigree rate can't be used for ages older than 5000 years (and all dates reported in this paper are clearly older) and, anyhow, the highly hypothetical molecular clock methodologies are anything but helpful when they become the main theme of a "research" paper.

I am not happy either because instead of using the already known SNP-based phylogeny of the haplogroup, they choose to treat the whole haplogroup as a single amorphous clade, when it is clearly structured. This isn't very helpful either.

Finally I am not happy either because they treat (again) the process of Neolithic spread in Europe as a single phenomenon, when it is in fact a complex array of various cultures, notably two different main vectors: one via the Morava-Danube and another via the Mediterranean coast. Both with origins not in Anatolia directly but in the Balcans. They ignore all these archaeological facts rather insultingly.

However for those who like to dig in the raw data, instead of just jumping to the too often biased and misleading conclusions, the paper still has some interest.

Notably I found figure 3 (haplotype structure) quite interesting. Here you have it with a crucial annotation for better understanding:

Click to expand

The crucial annotation is of course marking (with a dark red line) what is not part of R1b1b2a1, which is, as you can see a single branch of the star-like structure, but, unlike the others, is clearly not part of the fundamentally European haplogroup R1b1b2a1. Part of it is at the root of R1b1b2 and R1b1b2a but the rest of sub-branches must be derived, representing a distinct process centered in Turkey and nearby areas.

I dwelt on this matter in a previous post, so I'm not going to go all over it again here. Just for a quick reference a copy of the graph I posted then, showing major haplotypes and their relation with the various layers of the haplogroup:

Click to expand. Based on Alonso-2005.
DYS are 19-390-391-392-393.

As you can easily see most of the Turkish diversity belongs to the R1b1b2(xR1b1b2a1) part of the haplogroup structure. And sure I don't doubt that Anatolia or somewhere nearby is at the ultimate origin of R1b1b2. But there is a sharp distinction between that and what we find in Europe, which almost exclusively belongs to R1b1b2a1, a very specific sublineage.

And a sublineage that is very much ramified in a star-like structure, implying rapid demic expansion. When? That is not really the crucial issue as I see it. "Where?" should be the first question and a question that no paper has yet dealt with from the viewpoint of R1b1b2a1 on its own right.

In the past the lack of knowledge of the structure of the haplogroup may have served as excuse but not anymore. In fact any self-respecting geneticist should look at that SNP-based structure before dealing with STR-based haplotypes and take good notice of the distinctions.

And, if not, why not to include R1b1b1 (Central Asian) or even R1b1a (Italy and Africa mostly) and other R1b*? It is an arbitrary choice, poorly justified.

But, well, what do we get from this data set after we scrap off the extreme bias? For those who enjoy dealing with haplotypes in detail there is a long list in the supplementary material, which duly processed may provide very useful information.

I have not the time nor the resources to do that, so I have done something much simpler but also very informative: count the haplotypes by region as defined in figure 3 (above). Sadly France (incl. Basques), Germany, Netherlands, Denmark, England (incl. Cornwall) and Ireland are all dumped together in the category "other" ("West Europe" hereafter). Unlike the authors I do make the important distinction of what is R1b1b2a1 and what is not.

Follows the number of haplotypes by region and phylogenetic category (manual count so subject to minor error maybe):

R1b1b2(xR1b1b2a1):

• Turkey: 37
• West Europe: 20
• Iberia: 8
• Balcans: 2
• Italy: none

R1b1b2a1:

• West Europe: 198
• Iberia: 90
• Turkey: 13
• Italy: 11
• Balcans: 6

What does this say? That even between perfectly comparable regions such as Turkey, Italy and Iberia, the highest diversity for R1b1b2a1 is in the West. If you look again at figure 3 you'll notice that most Turkish haplotypes of this clade are derived from European ones, what implies back-migration after the formation and spread of R1b1b2a1, which must have happened in Western or Central Europe.

When did this happen? I am not sure but I have some things clear:

The structure of R1b1b2 does not correspond at all with what one would expect from a demic spread from the Balcans (not Anatolia directly) through two clearly distinct pathways, one to the Danube and Central Europe and the other through Italy to SE France and Iberia. Neither the Balcans nor Italy look particularly central nor we see two differentiated founder effects but only one.

Also the distribution of R1b1b2a1 in a cline that is, as the authors of this paper shamelessly admit, totally the inverse of what one could expect of a demic spread from SE Europe.

The structure of R1b1b2a1 in fact strongly suggests a spread from somewhere in the region described as "Other" and or Iberia, i.e. in Magdalenian Europe. This is in full agreement with the generally accepted theory that R1b1b2a1 spread from the Franco Cantabrian region after the Last Glacial Maximum, along with Magdalenian culture. It could have other explanations (Epipaleolithic flows, older Upper Paleolithic cultural dispersals like Gravettian or Aurignacian) but it just cannot fit within a Neolithic frame. No way!

How did it back-migrate to Anatolia? Possibly with the people who carried the rock art fashion to southern Turkey (Beldibi), which may have an offshoot also at Egypt, where some R1b1b2 is also found, as well as related haplogroup R1b1a. The exact process is still somewhat uncertain anyhow.

Sadly enough the authors have missed an opportunity to analyze the regional structure of this haplogroup in Europe. Hopefully someone else will eventually do it, helping to clarify the matter. The raw data is anyhow there for whoever wants to do it.


Update: a much more realistic geographic analysis of the diversity at the two different phylogenetic levels (by Aargiedude) can be found at this new post.

Lagar Velho child had modern human teeth

Mundo Neandertal leads me to a new research paper that shows that the teeth of the Lagar Velho child, arguably the most clear case of Neanderthal-Sapiens hybridization, in spite of being several thousand years more recent than the last known Neanderthals, had teeth that compare well with those of extant modern humans and, in particular, with young H. sapiens from La Madeleine (LM4).

Ref.: Priscilla Bayle et al. Dental maturational sequence and dental tissue proportions in the early Upper Paleolithic child from Abrigo do Lagar Velho, Portugal. PNAS 2010 (paywall or free access depending on your world region, supplementary material freely accessible in any case).

It is noticeable that more and more the support for the specimen being any type of hybrid seems to vanish. Computer Assisted Paleoanthropology clearly states that a comparative geometric-morphometric analysis of the Lagar Velho cranium clusters this individual with modern human children of age 3 to 4 and does not reveal Neanderthal affinities.

So I understand that is fair to say that at the present stage of research there is not a single specimen that can clearly be described as hybrid of H. sapiens and H. neanderthalensis, which is coincident with the growing genetic evidence against such hybridation.

Early European was mtDNA U2

A skeleton from the Markina Gora site (Kostenki XIV, Don basin, Russia) has seen its mtDNA sequenced by Svante Paabo's team, being classified as U2.

The researchers have developed a novel method that allows them to discern with apparent safety what is ancient DNA and what is contamination from modern handling. This method promises to revolutionize ancient DNA research.

Haplogroup U2 is relatively rare nowadays in Europe, being most common in South and Central Asia. Nevertheless it is also found across Europe at low frequencies and is believed to be old enough as for this finding to be consistent.

It is, of course, part of haplogroup U (in turn derived from macro-haplogroup R) and also part of the major subclade U2'3'4'7'8'9 (cf. PhyloTree) or, in other words, U(xU1,U5,U6).

Reconstruction of Markina Gora man by M. M. Gerasimov (source)

Markina Gora man is dated to c. 30,000 years ago.

Source: BBC News (a quite informative article).

Research paper: Johannes Krause et al., A Complete mtDNA Genome of an Early Modern Human from Kostenki, Russia. Current Biology, 2009 (paywall).

The AMH colonization of Europe

The colonization of Europe by anatomically modern humans (AMH, technically Homo sapiens) is generally accepted to be clear with the rapid Aurignacian expansion, c. 41-40 millennia ago (kya). But the real implications of the MP-UP transition are often controversial.

In this sense, John F. Hoffecker has made a nice review of the matter, to which I've been directed twice in the last weeks by Dienekes and Tim respectively, and that I feel it's about time to comment on here as well.

John F. Hoffecker, The spread of modern humans in Europe. PNAS, 2009.

The author reviews in few pages the available data on the MP-UP transition in Europe and their possible relations with each of the human species extant at the time.

In the conclusion, he argues for two waves of AMH colonization of Europe: first, beginning c. 48 kya, the Bohunician (that for him includes Bachokirian), that would originate in the Emirian culture of Levant, extending into the East Balcans, parts of Central Europe and even Eastern Europe. Second would be the proto-Aurignacian, originating in the Ahmarian culture (derived from Emirian) and extending specially southern European areas, like Italy and the Pyrenees. The Aurignacian proper would be a European culture, possibly originated in the Balcans (though others have argued Central Europe).

Let's review the sequence and groups in more detail:

Group A: would include the Szletian (Central Europe) and Chatelperronian (Western Europe) cultures. At least the later is clearly associated with Neanderthal remains and, by extension, many believe that Szletian would be too.

Group B: would include the Bohunician (SE, Central and Eastern Europe) and its possible ancestor the Emirian of the Near East (specially the sites of Bocher Tachtit in Palestine, Ksar Akil in Lebanon and Üçagizli in southern Turkey). The Bohunician dates to c. 48 kya (Bacho Kiro, Bulgaria) and the latest dates are of c. 40 kya. There are no clear forsenic remains associated to it but some indications of it being of AMH creation are the presence of ornaments in some sites and the relation between Emirian and Ahmarian, this one clearly product of H. sapiens.

Group C: would include the proto-Aurignacian of Bulgaria (Temnata) Italy, Pyrenees and other southern European sites (and even Kostenki 14 in Eastern Europe) and the Ahmarian of the Near East. This one is clearly the product of AMH and Hoffecker argues that some of the oldest H. sapiens remains of Europe (Pestera cu Oase and others in Rumania) could belong to this techno-cultural group, even if they are devoid of any direct artifactual associations. Some Italian proto-Aurignacian also shows personal ornaments and bone industry, which again suggest AMH manufacture. The proto-Aurignacian dates to c. 45-40 kya.

Other:

The Uluzzian of Italy (since c. 48 kya) again is not directly associated to any particular human species. For this reason many think it could be a Neanderthal creation. But Uluzzian shows again personal ornaments and bone industry and could therefore be associated to H. sapiens on mere cultural grounds.

The Kostenki culture of Eastern Europe (since c. 44 kya) is clearly of AMH creation and, while original, shows occasional similitudes with Italian proto-Aurignacian (Kostenki 14) and the later Aurignacian. It is rich in bone industry and includes the first eyed needles known.

The Aurignacian. Hoffecker suggests a Balcanic origin for this culture though it's best documented first in Central Europe (c. 40-38 kya), overlying the transitional industries. It is possible that its expansion was favored by the Campanian Ignimbrite eruption, the most catastrophic volcanic event in the area in some 200 milennia, which is dated to c. 40 kya. Guess it could be included in group C.
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European Upper Paleolithic (synthesis)

Just found a quite nice site in French on European Paleolithic, that may be of much help in revieweing European Prehistory, specially for the common reader without archaeological background. Authored by Thierry Koltes the site is Ma Préhistoire (my prehistory).

I'll borrow some maps from there for the purpose of this post. They may not be perfect but are good enough to illustrate European Upper Paleolithic:

1. Aurignacian.

Since c. 41,000 BP. This culture probably originated in West and/or Central Asia but finds its finished form only in Central Europe (Hungary), from where it expands rapidly westward into Moravia, Germany and the Franco-Cantabrian region, as well as Italy and other less populated regions (southern Iberia, southern Britain, some Eastern offshots), displacing and eventually driving the Neanderthals to extinction.

A more detailed map of Aurignacian sites (not perfect either) can be found here. See also this post.

Genetics: mtDNA haplogroup U seems old enough to have participated in this first colonization of Europe. Haplogroup U8a has been directly associated with it but U5 and U6, as well as other U subclades maybe, could well have been involved as well.

Anthropometry: the human type associated with Aurigancian is an archaic-looking dolicocephalic person like Combe-Capelle man.

2. Gravettian.

Since c. 30,000 BP. The origins are uncertain but again we found it evolved in Central Europe. From there it expands to the West and also to the East. It is the last pan-European culture. Areas like Italy or Eastern Europe will thereafter use Gravettian derived technologies.

Genetics: mtDNA haplogroup H (or its HV precursor) could perfectly have been involved in this second wave. Y-DNA R1b shows a similar star-like pattern and distribution, so I speculate that maybe R1 could have arrived then, spreading eastward as R1a and westward as R1b. Another possibility is that only R1b was involved in this phenomenon, with R1b1b2 and R1b1b1 heading respectively to East and West.

Anthropometry: the type associated with this wave is Cro-Magnon.

3. Solutrean.

Since c. 22,000 BP. Evolved in SW Europe, probably in Dordogne, and its main area is the Franco-Cantabrian region. In Mediterranean Iberia a local Gravetto-Solutrean evolves instead. While not shown in the map, a late Hungarian Solutrean is also known to have existed as well.

Solutrean peoples developed art and technology a lot. They had needles and fishing hooks, for example.

4. Magdalenian.

Since c. 17,000 BP, transitional forms since c. 19,000 BP.

The ultimate origins of the transition may have been in Germany, where Aurignacian pervivences are known to have existed till very late, but the transitional culture itself, known as Badegoulian is mainly from SW Europe.

Magdalenian replaced Solutrean in the Franco-Cantabrian region. Then it seems to have recolonized Central Europe, as the climate became slightly warmer after the last glacial maximum. A Magdalenian of late date is also found in southern Iberia - it has the peculiarity of lacking bone tools. It represents the cultural apogee of PaleolithicEurope, specially for its fascinating art.

Genetics: Many authors have argued that the Magdalenian recolonization of Central Europe represents the expansion of DNA markers from SW Europe, specifically Y-DNA R1b and mtDNA H clades, both of which show a clear star-like pattern suggesting a fast expansion.

Anthropometry: the Magdalenian human type has fully modern traits, even with impacted wisdom teeth (a byproduct of more narrow jaws). A good reference is Chancelade man.

Addendum.

The site does not follow up with Late Prehistory and I will leave it here as well. Just to mention a handful of issues:

Epipaleolithic cultures of Western and Northern Europe stem all from Magdalenian but experience a major reduction of size of tools, something also noticeable in other contexts of the same period (North Africa, West and South Asia). Cultures of SE and Eastern Europe instead stem mostly from Gravettian (epi-Gravettian cultures).

Mediterranean connections. Some groups in North Africa and West Asia have more or less clear apparent connections with European late UP cultures.

a. Anatolian Baldibian culture shows rock art that seems related with that of Europe, maybe reflecting a backflow through the Balcans.

b. The Epipaleolithic culture of the Zagros mountains, known as Zarzian, is derived from the Eastern European epi-Gravettian. Similarly to the Central European case, the Irano-Mesopotamian area seems to have been deserted in the LGM, and then recolonized, this time through the Caucasus.

c. North African Oranian (also Iberomaurusian) culture has been argued repeatedly to stem from some southern Iberian one (Solutreo-Gravettian?). The presence of Cromagnid types in North African late UP, as well as some genetic connections (mtDNA U6, H and V specially) may support this theory, that anyhow is contested by some who argue that Oranian is, like its Capsian successor, derived from cultures of Upper Egypt/Nubia. Nevertheless the strange "erratics" of Y-DNA R1b found both in Sudan and south of Lake Chad, together with the parallel African patterns of mtDNA U6, seem to suggest that the interactions and migrations were more complex and bidirectional.

d. Recently it has been discovered that late UP Upper Egyptians also practiced a realistic type of mural art that resembles that of Europe.

For a small collection of Paleolithic European and other skulls, see this post. For older general considerations on European UP, read this one.