Wednesday, May 10, 2017

DNA. "The Beaker Phenomenon" Olalde et al, 2017 (Update 15)

Here's a breakdown of "The Beaker Phenomenon and the Genomic Transformation of Northwestern Europe".  It's a pre-print at bioRxiv [here].

If you are new to this and don't really know much about the Beakerfolk or the European Neolithic/Bronze Age transition, I'll only say that this paper is possibly the most significant academic paper on this subject since it was first well-defined.

Grave 68/I1390, male - Haut-Rhin, France (L. Vergnaud), Antea Archéologie, "Current researches on Bell Beakers"  R1b1a1a2a1a2 - X2b4a (see more in update 10 below)


Abstract
Bell Beaker pottery spread across western and central Europe beginning around 2750 BCE before disappearing between 2200-1800 BCE. The mechanism of its expansion is a topic of long-standing debate, with support for both cultural diffusion and human migration. We present new genome-wide ancient DNA data from 170 Neolithic, Copper Age and Bronze Age Europeans, including 100 Beaker-associated individuals. In contrast to the Corded Ware Complex, which has previously been identified as arriving in central Europe following migration from the east, we observe limited genetic affinity between Iberian and central European Beaker Complex-associated individuals, and thus exclude migration as a significant mechanism of spread between these two regions. However, human migration did have an important role in the further dissemination of the Beaker Complex, which we document most clearly in Britain using data from 80 newly reported individuals dating to 3900-1200 BCE. British Neolithic farmers were genetically similar to contemporary populations in continental Europe and in particular to Neolithic Iberians, suggesting that a portion of the farmer ancestry in Britain came from the Mediterranean rather than the Danubian route of farming expansion. Beginning with the Beaker period, and continuing through the Bronze Age, all British individuals harboured high proportions of Steppe ancestry and were genetically closely related to Beaker-associated individuals from the Lower Rhine area. We use these observations to show that the spread of the Beaker Complex to Britain was mediated by migration from the continent that replaced >90% of Britain's Neolithic gene pool within a few hundred years, continuing the process that brought Steppe ancestry into central and northern Europe 400 years earlier.

(Update 1)
Reading now...

This might be helpful in the meantime.

Fig 3.  Bust and Boom in the Late Neolithic (Shennan et al, 2013)
Here's the blog link to the second paper by Timson et al, 2014

(Update 2)
  • Ok, so I was wrong.  Lactase Persistence is very low even in the British Bronze Age!!  What!
  • They absolutely, positively, unequivocally exclude admixture from Iberia (Neolithic ancestry of Beakers, they say is more similar to LBK populations (actually GAC and TRB).
  • Iberian Beakers have zero steppe admixture (except two girls in the North)
  • Dutch and British Beakers are identical
  • 95% of British Beakers are R1b P312.  However, this reduces in the MBA to 75%
  • 93.5% population replacement.  Wow again.  (Keep in mind this may be relative to immigration)
  • British Beakers were more vanilla and then became increasingly so. 
(Update 3)
"In central Europe, Steppe ancestry was widespread and we can exclude a substantial contribution from Iberian Beaker Complex-associated individuals, contradicting initial suggestions of gene flow between these groups based on analysis of mtDNA and dental morphology"
It'll be interesting to see how the Dulias paper approaches this problem.  H1, H3 and V in British individuals can't be Neolithic native, probably aren't Iberia (AFTER ALL), so what does this mean?

(Update 4)
  • R1b completely absent in Neolithic British samples n=20
  • I2a nearly completely absent in British EBA (which is admittedly biased for diagnostic BC)
  • No R1a in NE Scottish Beakers.
(Update 5)

I'm looking at the mtdna of Beaker individuals.  Excluding the questions traditionally circling H1, H3 and V, a lot of the rest of it looks surprisingly steppe-like.

(Update 6)

After reading through the first time and looking at the Balkans paper, several things ranging from 1) effectively proven to 2) more likely with new factual weight being added, now seem possible:
  • Beaker appears to have come directly from the Pontic-Caspian steppe while absorbing LBK-like or admixed ancestry. Census estimates and other data supports this. 
  • Many of their mito-profiles look steppic.
  • The Ukraine looks mixed
  • Skin, eyes and LP basically identical to Yamnaya, changes over time. (Which I still don't get)
  • Not a trickle of people.  Waves of immigrants first into Europe, then the isles with massive population increases following (older papers below).
  • Areas of Europe less affected initially get theirs in the Bronze Age.
  • U106 ~= Veluwe Beakers
(Update 7)

Looking a little closer at the Iberian remains/contexts associated with Beaker culture.

First, it should be understood that the remains and contexts are what they are.  Plus, even with messy and uncertain contexts, certain ancestry would or wouldn't be there, so this is not a criticism for including these particular remains.  I'm bit surprised at how loose the connections seem between people who look like otherwise ordinary Neolithic Iberians and Beaker associated artifacts.

In other words, these are mostly collective tombs and caves with complex and disturbed deposits. 
Here's a few of those and the identifications, the first two bolded are R1b with no steppe admixture, the second two are girls with steppe admixture)

-  Galeria da cisterna, tomb (I0839 and I0840)
-  Cova da Moura, cave (I4229)
-  Paris Steet, Barcelona (I0257, I0258, I0260, I0261, I0262, I0263, I0823, I0825, I0826 and I1553)
-  Arroyal I, Burgos (tomb Roy5 (I0462) girl with first steppe admixture in Iberia.  Also (I0458, I0459, I0460, I0461)
-  Camino de las Yeseras (a flat head woman I4245)
-  I4247 (collective inhumation)
- Camino del Molino, Caravaca de la Cruz (I0453)

The Olalde authors made a compelling case that the source of Neolithic admixture in Beakers was largely LBK-based (simply meaning Northern Europe - and actually more close to a mix of Globular Amphora and Funnelbeaker ancestry which may be more directly indicative of significant Corded Ware ancestry of Northern Europe).  Very compelling.  However, I am not really sure that a single individual, other than the woman from Camino de las Yeseras, was actually a truly diagnostic Beaker.

I'm not saying this picture will change much with more testing, but I do believe steppe ancestry is in Iberia and it may be more elusive.

(Update 8)

The authors confirm that the approach to understanding mobility through isotopes is that first generation movers are always going to be few and far between, even when ~93% of a population is immigrant descended.  Valuable data, just needs interpretation beyond turtle-level cautious.

(Update 9)

This is interesting.  The burial of  "Rue de Phaffenheim" is of a 30 year old woman (I1391) who was buried in the supine position with legs flexed right, very similar in format to a Yamana burial, although she is from the mature Beaker phase.

A fascinating aspect of her vestment is that many v-perforated buttons dot the right hemithorax.  In other words, she wore what appears to have been a right-buttoning sweater, coat or blouse.  Before the modern era, the gender-differentiation of button and belts may have been like men for both sexes, because we are right handed.  So this may be something to look into later...

(Update 10)

I added a picture and a source on the Archer from France (I1390).  He's also the very first burial on the burials page at the top of the blog "30 Beaker Burials".  He was buried with a quiver of arrows.  Genetic testing proved that a young man (I1389) buried near him was a first degree relative, probably his brother, since they both have also have the same mtdna X2b4a.

In fact, the radiocarbon dates may suggest they were closer in age in life, than the time difference in death. (Lines 270 and 271 in Supp. 1)

(Update 11)

La Fare (Forcalquier, Alpes-de-Haute-Provence) (photo Lemercier) S14/I2575  Y-unsucessful? mtdna - K1c1
Notice the copper knife behind the ear of this French Bell Beaker of the Rhone-Ouvèze group.  He was about 5.6 feet or 1.72 meters tall and a "so-called "alpine" cranial architectural type".  In other words, he had a big, weird Beaker-head.  More pictures on Burials page 2.
Here's the text:  "Le Campaniforme dans le sud-est de la France"

(Update 12)

The Italian Beaker from Parma may be described in this document [here] I believe.  He was (I2478) R1b1a1a2a1a2 - K1a2a.  Other men were present and fairly seasoned 50's and 60's of age.

(Update 13)

Another unusual situation where a man has been buried as a woman.  You'll remember previously a similar situation in the Allentoft paper concerning Kněževes - grave 8 who was a woman buried as a man, with man gear (genetically confirmed a woman).  This has been reported in the CWC as well.

In this case, the man in Grave 1 (line 675 of the Supp. 1) was buried in the female, head-south format with a copper awl, something only Beaker women have been buried with.  The other individuals of the cemetery were buried according to normal rites.

Since they did not appear to DNA test grave 1, I'm not sure if the archaeologist's determination was based on skeletal metrics or confirmed through DNA and not published.  I'd be cautious in projecting too much here.  There's a lot of weird stuff to explore with regard to graves 1) cosmology 2) coming of age 3) married vs. non-married 4) honored and dishonored 5) priestly people and interestingly, 6) noncommunicants, it seems at times.

(Update 14)

When looking closer at the Beaker y-chromosomes, the haplogroup frequency becomes more extreme.   Here's what I mean:

(I1767) is a young male that is the only British male associated with a non-R1b haplotype in the Beaker period from this study.  He's I2a2a1a1a.  Besides his radiocarbon date, his grave contains no diagnostics other than he was a single burial, and he and a female were not in a gender direction (which doesn't necessarily mean anything in Britain).  Not really relevant but interesting, he was intentionally mummified.

(I2364) was a male from a double grave, head south on the western bank of the Danube around Budapest, Hungary.  While he's buried in a large cemetery that contains Beakers, only a small fraction of those graves actually have diagnostic goods, his didn't.  He was Y-chromosome H2.

That leaves two more Hungarian Beakers: (I2786) who was definitely a Bell Beaker having I2a2a, and (I3528) who was probably so G2a2a1a2a1.  One German Beaker (E09538) G2a2a1a2a1a.

I'm totally excluding the Iberian collective contexts where 8 y-chromosomes were published.  I'm just not totally confident that these folks represent people who were ever associated with Beaker cultural materials.  Totally shooting from the hip, but a number as extreme a 3/42 BBC males being associated with non-R1b male lines seems possible.

So what is that like anyway, 0.7%?  But keep in mind that the non-Iberian Beakers all had steppe admixture, so that would raise that number a little with a more reasonable criteria. Plus, these are just Beakers, not all Western Europeans.

(Update 15)

I'm surprised and intrigued by the absence of Corded Ware Culture R1a lineages, especially in Scotland, the Netherlands and Germany.  This presents a whopping dilemma.  It would seem very reasonable to believe that BBC had CWC ancestry, but I'm not sure that math works.



As soon as the genomes are available, we'll see some improved or alternate models.  Also, there's more Beaker genomes coming!


"The Beaker Phenomenon And The Genomic Transformation Of Northwest Europe"

Iñigo Olalde, Selina Brace, Morten E. Allentoft, Ian Armit, Kristian Kristiansen, Nadin Rohland, Swapan Mallick, Thomas Booth, Anna Szécsényi-Nagy, Alissa Mittnik, Eveline Altena, Mark Lipson, Iosif Lazaridis, Nick J. Patterson, Nasreen Broomandkhoshbacht, Yoan Diekmann, Zuzana Faltyskova, Daniel M. Fernandes, Matthew Ferry, Eadaoin Harney, Peter de Knijff, Megan Michel, Jonas Oppenheimer, Kristin Stewardson, Alistair Barclay, Kurt W. Alt, Azucena Avilés Fernández, Eszter Bánffy, Maria Bernabò-Brea, David Billoin, Concepción Blasco, Clive Bonsall, Laura Bonsall, Tim Allen, Lindsey Büster, Sophie Carver, Laura Castells Navarro, Oliver Edward Craig, Gordon T. Cook, Barry Cunliffe, Anthony Denaire, Kirsten Egging Dinwiddy, Natasha Dodwell, Michal Ernée, Christopher Evans, Milan Kuchařík, Joan Francès Farré, Harry Fokkens, Chris Fowler, Michiel Gazenbeek, Rafael Garrido Pena, María Haber-Uriarte, Elżbieta Haduch, Gill Hey, Nick Jowett, Timothy Knowles, Ken Massy, Saskia Pfrengle, Philippe Lefranc, Olivier Lemercier, Arnaud Lefebvre, Joaquín Lomba Maurandi, Tona Majó, Jacqueline I. McKinley, Kathleen McSweeney, Mende Balázs Gusztáv, Alessandra Modi, Gabriella Kulcsár, Viktória Kiss, András Czene, Róbert Patay, Anna Endródi, Kitti Köhler, Tamás Hajdu, João Luís Cardoso, Corina Liesau, Michael Parker Pearson, Piotr Włodarczak, T. Douglas Price, Pilar Prieto, Pierre-Jérôme Rey, Patricia Ríos, Roberto Risch, Manuel A. Rojo Guerra, Aurore Schmitt, Joël Serralongue, Ana Maria Silva, Václav Smrčka, Luc Vergnaud, João Zilhão, David Caramelli, Thomas Higham, Volker Heyd, Alison Sheridan, Karl-Göran Sjögren, Mark G. Thomas, Philipp W. Stockhammer, Ron Pinhasi, Johannes Krause, Wolfgang Haak, Ian Barnes, Carles Lalueza-Fox, David Reich


23 comments:

  1. I'll read the article later, but this particular statement bothers me:

    " we observe limited genetic affinity between Iberian and central European Beaker Complex-associated individuals, and thus exclude migration as a significant mechanism of spread between these two regions..."

    This seems to define migration as necessarily a wholesale replacement of populations, as in the British colonization of North America. But obviously, any claim has to account for the distribution of the bell beaker design. This sort of non-migration claim assumes a movement of ideas that never gets explained in the details. Can we imagine that the bell shape and surface design elements moved from village to village, from Iberia to Germany? I think it's far more plausible to assume that people did the traveling, and their pots - or their beaker design preferences - traveled with them. It could have been a small number of people, but it would have to be enough to impress the natives and get them copying the new folk. If you don't call that migration, there needs to be a new term to account for it.

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    1. I'll have to dig in to the details of the Iberians and how they were buried.

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    2. Traders, could explain it, I suppose. I know that making comparisons to cultures that existed much later in history might be a wrong approach, but Pheonicians also traded along the Atlantic, despite there being no colonies in Europe west of the Guadiana river in SW Iberia, their influece was still felt in places they never actually settled - and where, expectedly, they left no genetic trace. Could something similar have happened in NW Europe with early Iberian Beaker folk?

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  2. Not entirely related to this, but you might want to check the paper "The Population Genomics Of Archaeological Transition In West Iberia" at http://biorxiv.org/content/early/2017/05/10/134254

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  3. Hey BBB,

    what do you make of R1b1 and R1b1a (xL23) in the Barcelona beakers? Also, as per the supp. material it seems the oldest Beaker with R1b-L53 is found in Haute-Savoie with relatively minor steppe aDNA.

    Am I being stupid here or does the data contradict the authors' interpretation?

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    1. I hadn't noticed that. Even relatively minor can be significant that early due to local admixture. But as I've mentioned before Iberia may be more mixed than these 20 samples show.

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    2. I agree, though the two R1b1 Beakers from Barcelona completely lack the steppe signal. The Hégenheim woman in eastern France is identical to the Spanish beakers, though later eastern admixture becomes widespread in these regions. It seems France is where the East-West admixture initially took place.

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    3. Also MBA southern Portuguese samples have no CHG. They happen to be R1b P312 though, and the paper say they have steppe ancestry. I wonder what they mean by 'steppe' if CHG isn't present

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  4. After letting this stuff sink in a bit, the most burning question in my mind is: "Why and in what manner did the Steppe folk in Central Europe decide to adopt so much of the Bell Beaker material culture?"

    Pretty much all of the narratives I've seen have involved outsider steppe-genetic people with superior technology/culture/whatever emulated by less technologically advanced first wave European farmers. This scenario is more like the Roman invaders of Greece adopting Greek culture instead of their own. The details matter of lot for purposes of historical linguistics and in general for understanding who two demographically and genetically almost completely disjoint populations end up both being part of an Eneolithic/Bronze Age superculture with a huge geographic scope.

    What was the pitch made by the Iberian missionary/traders? Bronze?

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    1. Well, if they are able to get another 80 Iberian burials, preferably not in dolmens or caves, but actual single burials or elite VNSP burials, the steppe ancestry may be there after all.
      Iberia is much more developed than Holland or England at this time, I bet the steppe ancestry is there, maybe diluted in restricted to certain family groups. I just don't think these collective tombs are necessarily informative unless you already had the assumption that Iberian Beaker meant native Neolithic Iberian, which I never believed.

      Also, the strangely early AOC beakers in Iberia and the apparent need for the Portuguese castles doesn't suggest to me there was two distinct Beaker worlds.

      I haven't read the Western Iberian Bronze Age paper yet, but I think it offers a clue to the presence of Continental-like Beakers in Iberia that just aren't well represented in the megaliths of this paper.

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    2. Hoh hum ... I see an analogy in the current technologically superior USA and the adoption by significant portions of the population and legislators of creationism ... (If we can explain that of today, maybe we can explain something similar from thousands of years ago.)

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  5. "Valuable data, just needs interpretation beyond turtle-level cautious."

    Surely not cat-level cautious, however, because that could be deadly.

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  6. Trade routes have been there since time immemorial, trading crops and seeds, stone cores, news ... Later metals, glas, more news, amber, slaves prehaps. Or embassies offering hostages to settle a deal. Marriage brokering. Exchange of foster-children by elites (cf two steppe-admixed girls in Northern Spain?) It seems Maykop was a centre in an intricate web of trade-routes. When the steppe emptied (for whatever reason) they didn´t just bugger off into the blue, I reckon. Also it seems that the Portuguese trading-places along the Atlantic sea-routes needed to be protected against competition, whoever they were. Originally I imagine most trade was water-bourne. But then the mobility on the steppe (horses, oxen, carts) changed the pace and the scope of trade and politics. New elites formed, and the egalitarian matriarchal farmer-communities were either recruited, absorbed, bypassed, and/or possibly occasionally wiped out. By 1200 BC everybody knew about everybody, and the battles that were fought probably involved complicated coalitions from far and wide.
    I am disinclined to imagine the Yamnaya as whooping and hollering savages on horse-back, but would rather cast them as diplomats. Calculating, but not without codes. Pure speculation though, but I think the ´murdering hordes from the steppe´ were a later thing. Eneolithic and BA must have been the advent of high culture, and elitist, paternal social structure where prestige (and honour, of all kinds) was everything.
    I apologise if all of this is a bit of an open door being kicked in, but the idea of ´globalisation´ comes to mind. Or am I just stating the obvious?

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    1. I agree with everything you have said. Patriarchy is undersold as being a driving factor in a number of respects. That evolved into the idea of a landed male citizenship that was so important among Greeks and Saxons. So murder and mayhem, maybe more in the LBA, but much of the change internal social pressure.

      Also the proximity of Beaker settlements to other cultures and the exchange of influences is an indication of those relations as well.

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    2. "By 1200 BC everybody knew about everybody, and the battles that were fought probably involved complicated coalitions from far and wide."

      I think this is a great point, and it's supported by the surprising discovery of the massive battle at Tollense dated approximately 1250 b.c. Apparently there were warriors there from all over Europe: http://www.sciencemag.org/news/2016/03/slaughter-bridge-uncovering-colossal-bronze-age-battle

      I'd love to see a DNA study of the remains found at the battle site.

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  7. @Dan May: Some of them must have been sequenced, or how would I have known that they were from all over Europe? Just don't remember: I'm lazy and old. I remember vaguely I used to remember every little detail ... I also remember speculations that mercenaries might have been involved, professional soldiering! Some horses were there, dead ones! May not have done much good ... Bit like Flemish peasants beating French poofters in shiny armour(?). At some point dreamers will always get a horrible spanking from reality, but that is my projection: maybe the pros won, at some cost. Somebody got spanked for sure!
    @bellbeakerblogger: Thanks! Wondering about where the plus-value was for patriarchy. And not being a sexist male patriot, who's appreciative of the female element, I still speculate that in them days there was more heart in patriarchy. I know about football hooligans and creationists and people who need three or more mistresses in as many Paris apartments (at the cost of the simpletons who put their trust in them), but this must be the pendulum swinging. I suspect many BA male sexists were of a very noble and deep sort. People who knew how to keep contracts, honoured friendship, and generally tended to look beyond the material benefits. Can I take one more salmon leap here?
    Islamic suppression of women is an expression of a deep fear, which by now has outlived its cause. It's horrible and degrading, no question! It's high time these people began to understand what it is that they're passing on to their children! But in times gone by, probably well into the times of the prophet, this fear did have some substance. I'm now thinking of the crazy risks young Minoan males were taking in these bull-leaping contests. I'm thinking of the well-manicured young males - "with not a blemish on their bodies" - we keep finding in the bogs of Northern Europe. "Dar" in Dutch means a male bee, the one who gets his wings gnawed off after mating with the queen (Or was that ants? Mixed up old-guy-stuff!), but it is also an old word for 'king' ('Dare', like in Curoi McDare). Cyrus son of Darius. Every year there was a new king in some Irish annal (forget which manuscript. Dun cow?). It's difficult to assess, for of course the scribes who produced their versions of the old legends had an agenda of their own: they hang onto some things out of respect, but discarded other things, not to mention what they may have invented. When Irish scribes invented, they changed their sources somewhat, correcting imperfections according to the 'truth'. All in all I suspect that at one point in our history promising, nice, young, attractive, athletic males were selected for the honour of being garotted and dumped in a swamp, along with so many of other nice things, such as swords, gold adornments, etc (no torcs?). Given back to the earth: nice! And not so nice ...!
    The females are the survivors, the males are expendible. I almost agree here! Still a male fear of female power does make some sense. I don't want to discuss female infidelity as a lever on men, another source of fear. Don't want to talk about myself!
    DOING!! Time's up: haven't been able to summarise Proust in 10 seconds!

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  8. PS: So I was merely stating the obvious. I'm OK with that!

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  9. Well I thought there might be an mtDNA H6a1a found in BBC. And yes, West Frisia Netherlands had a H6a1a in a well used barrow. And today I have an 1 exact match in the Netherlands, 2 in England, 3 in Ireland, 1 in Serbia, 1 in the US and 4 in Germany(my last known maternal Grandmother left Northern Bavaria in 1850) So this looks like a CWC Steppe ancestry females(with ancestors in the PC Steppe Cultures) joining the BBC. What stories they could tell!

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  10. @ BBB

    "Also, there's more Beaker genomes coming!"

    Do you know something about future papers about Bell Beaker samples?

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    1. There's some Moroccan Beakers at Stanford. Plus there's a few that have been tested, but still no results

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  11. What does "British Beakers were more vanilla" mean? Vanilla as in colour? vanilla as in plain or uninteresting? Thanks

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    1. They may have been very plain and uninteresting ironically, but here I'm referring to coloration traits compared to the Neolithics

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