R1B & Bell Beaker

markesmith

R1b and its subclades seem by far and away the dominant haplotype in Ireland (and Wales). Recent research is saying that R1b arrived about the time of the Beaker people. Did they really take over the island so comprehensively?

Ipso

Just because it is the dominant haplogroup now doesn't mean it was back then.
In Ireland the high rate seems to be the result of Genghis Khan effects from about 500 AD onwards, where you had powerful chieftans having a lot of children with various women and then the male children doing the same.

I think in Europe the big overall genetic differences are on a North to South basis, and R1b spread from east to west.
That may be an indication that it's spread was more of a male dominated elite group.
Dubhthach will be able to explain better.

dubhthach

Well it depends on what variant of R1b you are talking about. There's some debate that "Bell Beaker" material culture may be linked to R1b-P312 and subclades, for example the remains tested from Germany came back as been R1b-M269+, R1b-U106- (they didn't test for R1b-P312).

In context of European R1b the major clade is basically M269 -> L11

Under L11 the two big branches are P312 and U106.

So far none of the ancient-DNA recovered form mesolithic and Neolithic remains are R1b they all show up as either Haplogroup G (like Oetzi the iceman) or Haplogroup I. Even more interesting the 8,000 year old remains from Northern Spain were Haplogroup C!

Dr. Michael Hammer showed the following diagram for his proposal on expansion of R1b into western Europe in the post-neolithic era.



As an aside I came across the following map which is interesting (for example it uses DF27 which is upstream of Z195 on Hammer's map)



What's evident on studies of distrubition is that the various R1b clades in the west are younger, the flow of SNP's appear's to be from east to west. With highest diversity found in the east. Not really surprising given that the oldest known Haplogroup R* (parent of R1(a,b) and R2) dates from 25,000 year old remains from Siberia.

dubhthach

Anyway the key issue in Ireland as to why some lineages multiplied to such high levels is probably due to the structure of Gaelic society before it's destruction by the Tudors and the Stewarts.

Here's an extract from Nicholls seminal "Gaelic and Gaelicized Ireland during the middle Ages"

---
One of the most important phenomena in a clan-based society is that of expansion from the top downwards. The seventeenth-century Irish scholar and genealogist Dualtagh Mac Firbisigh remarked that 'as the sons and families of the rulers multiplied, so their subjects and followers were squeezed out and withered away; and this phenomenon, the expansion of the ruling or dominant stocks at the expense of the remainder, is a normal feature in societies of this type. It has been observed of the modern Basotho of South Africa that 'there is a constant displacement of commoners by royals [i.e. members of the royal clan] and of collateral royals by the direct descendants of the ruling prince;, and this could have been said without adaptation , of any important Gaelic or Gaelicized lordship of late medieval Ireland.

In Fermanagh, for example the kingship of the Maguires began only with the accession of Donn Mór in 1282 and the ramification of the family - with the exception of one or two small and territorially unimportant septs - began with the sons of the same man. the spread of his descendants can be seen by the genealogical tract called Geinelaighe Fhearmanach; by 1607 they must have been in the possession of at least three-quarters of the total soil of Fermanagh, having displaced or reduced the clans which had previously held it. The rate which an Irish clan could itself must not be underestimated. Turlough an fhíona O'Donnell, lord of Tirconnell (d. 1423) had eighteen sons (by ten different women) and fifty-nine grandsons in the male line. Mulmora O'Reilly, the lord of East Brefny, who died in 1566, had at least fifty-eight O'Reilly grandsons.
Philip Maguire, lord of Fermanagh (d. 1395) had twenty sons by eight mothers, and we know of at least fifty grandsons. Oliver Burke of Tirawley (two of whose became Lower Mac William although he himself had never held that position) left at least thirty-eight grandsons in the male line.

Irish law drew no distinction in matters of inheritance between the legitimate and the illegitimate and permitted the affiliation of children by their mother's declaration (see Chapter 4), and the general sexual permissiveness of medieval Irish society must have allowed a rate of multiplication approaching that which is permitted by the polygyny practised in, for instance, the clan societies of southern Africa already cited.
---

(the above is my transcription so excuse any spelling mistakes)

markesmith

Thanks for this dubhthach, excellent explanation as always.

However, I find the DNA side of things confusing, and after reading Oppenheimer's book and then finding much of it had been discredited, I'm confused about the whole area.

Flow of R1b from east to west - would that tally with the corded ware culture (a precursor to the Beaker Culture), which spread from the east?

Also, the east-west spread of R1b - would that explain why it was mistakenly thought of as the 'Neolithic farmer' gene, in that it spread from the (middle) east?

Finally, what are your thoughts on the intrusion of Beaker culture as being the agent of proto-Celtic languages into Ireland?

dubhthach

My understanding of Corded Ware (which is slight to be honest) is that geographically it's east/north-east of the areas of highest Beaker activity. There's obvious overlap within what is now Germany.

Maps taken from wikipedia (so no idea on accuracy)



Bell beaker:



There is some debate about Beaker whether it's a material-culture package spread by trade/contact. There's a possibility of that, after all the oldest beaker material appears in Iberia (if memory serves me correct). At the moment the minimum that we can say is that R1b has been found in Beaker remains from Germany, there's no ancient DNA from Iberian beakers so we can't really tell.

Eupedia has the following map for the movement and expansion of various R1b lineages.


What's evident that within European R1b there is a major division under R1b-L11 into two clades called R1b-U106 and R1b-P312 (there's a new smaller clade called R1b-DF100 -- this probably makes up bulk of R1b-L11* men in Busby/Myers). Anyways both P312 and U106 date from the Bronze age (they are on order of a minimum of 4,000+ years old)

Their current distrubitions look like the following
P312 (unfortunatley labeled "Italo-Celtic" on Eupedia site)


U106 (S21)


Anyways the key thing will be ancient-DNA, all of those maps are based on current distribution. On current Indo-European languages there does seem to be a spilt between R1a and R1b which is interesting as the two lineages probably spilt on order of 15-20k years ago. Some have proposed that R1a is the "Indo-European lineage" (to be honest it's probable ye had multiple lineages) and that say for example "R1b heavy group" was assimilated in. It's all speculation though really until we get lots of detailed ancient-DNA.

As for Proto-Celtic and Beakers. Well it's probable that Proto-Celtic is a language of the Bronze age (Greek and Hittite were both differentiated circa 1500BC), I personally think Beakers are too early, though perhaps you had some form of "early western Proto-Indo-European dialects".

The major P312 subclades such as L21, U152, DF27 are all on order of 3700-4200 years old. The highest diversity of L21 appears to be on the continent, given that 70% of Irishmen are L21+ this points to mass replacement of male lineages in Ireland, by one lineage that dates to middle Bronze age at the earliest and probably arose in what is now basically France. (Basically all L21+ men are descended from the first man born carrying L21 snp).

Regarding the "neolithic" I believe a number of studies in period 2009-2011 pointed to R1b been linked to Neolithic, these were based off a calculated "mutation rate". Part of the problem in sense is that none of the neolithic ancient-DNA has come back as R1b.

Ipso

Isn't there a bell beaker site in Ireland (maybe Ross Island in Kerry) that is much older than others in the Isles?

dubhthach

Not sure about wether it's older then elsewhere in the isles but here's an article I came across on academia.edu about Ross Island (if memory serves me right my Dad might have a book about it at home)

Ross Island and the Origins of Copper Metallurgy in Ireland.

http://www.academia.edu/4809361/Ross_Island_and_the_Origins_of_Copper_Metallurgy_in_Ireland

What's important from what I recall is that Ireland had one of the big sources of Copper in western Europe at the time, so if the site is earlier then elsewhere in the isles it's due to importance of Copper to specific material culture.

dubhthach

With regards to disconnect between Neolithic and modern populations the following has just been published:

Tracing the genetic origin of Europe's first farmers reveals insights into their social organization
http://biorxiv.org/content/early/2014/09/03/008664

Dienekes has article about it here:
http://dienekes.blogspot.ie/2014/09/y-chromosomes-and-mtdna-of-early.html

Surprisingly, Y chromosome haplogroups, such as E1b1b1 (M35), E1b1b1a1 (M78), E1b1b1b2a (M123), J2 (M172), J1 (M267), and R1b1a2 (M269), which were claimed to be associated with the Neolithic expansion [23–25], have not been found so far in the 6th millennium BC of the Carpathian Basin and Central Europe. Intriguingly, R1a and R1b, which represent the most frequent European Y chromosome haplogroups today, have been reported from cultures that emerged in Central Europe during the 3rd/2nd millennium BC, while a basal R type has been reported from a Palaeolithic sample in Siberia [60] in agreement with a proposed Central Asian/Siberian origin of this lineage. In contrast, G2a has not been detected yet in late Neolithic cultures [42,43]. This suggests further demographic events in later Neolithic or post-Neolithic periods.

Considering the entire set of 32 published NRY records available for Neolithic Europe thus far, the low paternal diversity is indeed quite remarkable: G2a is the prevailing haplogroup in the Central European and Carpathian Basin Neolithic, and in French and Iberian Neolithic datasets [36,40,41]. There are only two exceptions, namely one E1b1b (V13) [41] individual from the Avellaner cave in Spain (~5,000-4,500 BC), and two I2a [40] individuals from Treilles, France (~3,000 BC).

markesmith

dubhthach, any way you can do an ELI5 on these latest findings?

I'm having a hard time...comprehending what this means. Sorry!

dubhthach

ELI5? sorry don't follow, also which findings? The ones in this thread or something newer? (Corded Ware / ANE that's in the news at moment?)

cfuserkildare

Hey folks,

Excuse my ignorance since DNA related subjects would not be my best subject,

But,

Unless we are dealing with actual physical samples, is not most of this theoretical?

Surely we don't actually know when a race reached Ireland unless we have viable DNA samples since items like pottery could well have been copied from pre-existing cultures?
The existence of Corded Ware as an example could mean that the spread of a particular culture needn't be factual, but that 2 cultures could have say met or lived near without actually being the same culture?

I know I am wandering a bit, trying to formulate theory in my head as I type it.

dubhthach

It's a good quetion, I'm gonna give a long answer (which may confuse things even further )

In case of Ireland there are as let no published ancient-DNA. As a result we have to infer what the situation was based on two factors.

1. Modern population genetics of the Irish population compared to other European populations (for example Scottish, English, French, Spanish, Germans, Scandinavians)
2. Published ancient-DNA results from continental Europe

The published ancient-DNA at the moment covers Mesolithic (Spain, Sweden, Luxemburg), transition populations (hunter gathers with contact with farmers) from Sweden, Neolithic populations from Iberia, France, Germany (Stuttgart) and Scandinavia. Some ancient DNA from Bell Beaker in Germany.

There is a major publication with regards to corded ware ancient-DNA expected in next while, it hasn't been released let.

What's evident is that modern populations are "shifted" from both Mesolithic and Neolithic populations. I would think a good example is basically "Latin America" if you compare pre-columbian contact populations in the likes of Mexico with modern Mexican population, then the modern population is "shifted" towards European/African populations. What you see is that most Mexican's have some level of Mesoamerican (Amerindian) ancestry, this can vary from 5% all the way up to 95%+, with the average Mexican propbably having between 30-60% amerindian ancestry. The rest of their genome is made up of European and African admixture.

What we see in Europe is exactly the same process (Admixture) except instead of acting on a 4-500 year period it's occurred over the last 8,000 years.

The result is that at the most basic that all Europeans are descended from the "blending" of three distinct population groups, namely:

• Western Hunter Gathers (WHG)
• EEF - Early European Farmer
• ANE - Ancient North Eurasian

Samples from the Mesolithic basically fall into WHG group, whereas EEF consists of a mixed population made up of mixed WHG ancestry with that of a group called "Basal Eurasian" (which had spilt from other Eurasian populations at a very early point on trip out of Africa). Basically farmers show some affinity to populations now found in Eastern Mediterranean.

Subsuqently a third ancestral population was added into the mix probably in the late neolithic this has been identified with a population that at originated at one stage in what is now Siberia. The 25,000 year old siberian boy whose genome was sequenced last year belong to it. This population contrubited to both modern Europeans as well as to Amerindians (mix of it and a East Asian population). What's interesting about the 25k year old boy (MA-1) is that his Y-Chromosome belonged to Haplogroup R* (eg. he was negative for R1 and thus R1a or R1b).

Different modern European populations have varying levels of these "three ancestral components". So for example in the recent study published:

Scottish sample:

• WHG: 42.8%
• EEF: 39%
• ANE: 18.2%

French sample:

• WHG: 31.1%
• EEF: 55.4%
• ANE: 13.5%

Greek sample:

• WHG: 5.8%
• EEF: 79.2%
• ANE: 15.1%

Anyways if we leave aside the look at the whole genome (above with three samples) we see that there appears to be a near total turnover of male lineages based on what we know of ancient DNA.

The oldest western European male so far sampled is an 8,000 year old hunter gather from Northern Spain. He belonged to Haplogroup C (found at very low levels in modern European men, also found in Aboriginals). Other mesolithic male samples belong to Haplogroup I, whereas Neolithic appears dominated by Haplogroup G (Otzi the iceman, neolithic cemetry in Southern France). What's important about this is we so far see a complete lack of Haplogroup R1b in western European ancient-DNA from before the bell beaker era.

In context of Ireland this is important as Busby's study (published by Royal Society) estimates that 90% of current Irish male population belongs to Haplogroup R1b. So using neighbouring European countries as a proxy it's fairly certain that there were no male's of Haplogroup R1b in Ireland during either the mesolithic or early Neolithic.

There's some debate that R1b in western Europe spread with the beaker culture, again until we get a couple dozen samples of viable male ancient-DNA from beaker remains this is sorta conjecture. We do have the beaker sample from late Neolithic Germany that came back as R1b, this is oldest known sample of R1b let found.

(see: Lee, E. et al. (2012), Emerging genetic patterns of the European neolithic: Perspectives from a late neolithic bell beaker burial site in Germany, American Journal of Physical Anthropology, online 3 May 2012 ahead of print.)

What's evident with R1b in Europe is that even though it increases as you go westward (90% in Ireland) it's diversity actually drops, what's evident is that the more diverse upstream subclades are found as you go eastwards, this points towards an expansion westward.

Dr. Michael Hammer (Univeristy of Arizona, Familytreedna) presented the following slide at the FTDNA (Familytree DNA) conference last year, with regards to R1b showing this, the clades dominant in the West are actually more recent/less diverse:



The prime example been R1b-M222, which Trinity College linked to the Uí Néill and the Connachta of medieval Ireland. This haplogroup probably dates to the Iron age at the earliest let it makes up almost 12-15% of male population in the likes of Donegal!

That one lineage could reach such a high level in a short period reflects probably societal structure in Gaelic Ireland.

Anyways back to your question about "Race" and two cultures adopting elements of each other's cultural package and not intermixing, well on first part sure. If my neighbour has a better way of making a pot I'll probably copy it. However what we are seeing isn't necessary one "race" coming in and taking over, instead we are seeing blending process.

A new group would come in with some improved or superior technological population and over time it would eventually merge with the proceeding groups through intermarriage. So for example in Scandinavia, modern Swedes appear to be mix of "Hunter gatherers" and Farmers (who show affinities with modern mediterranean populations such as Sardinians). They inherited from both, they are closer to the hunter gatherer population but have been shifted by the influx of early farmers.



see: (Skoglund 2012, Origins and Genetic Legacy of Neolithic Farmers and Hunter-Gatherers in Europe -- Science 27 April 2012: Vol. 336 no. 6080 pp. 466-469 DOI: 10.1126/science.1216304 )

cfuserkildare

Interesting,

As I said before, my knowledge would be more hands on practical as opposed to Genetics and DNA.

I will need to do some more research before I can get to grips with this properly.

Cheers dubhthach.

Wibbs

As always Dubhthach when you post you bring real value and fantastic scholarship and on top of that make it all the clearer for the rest of us. Always a true treat to read.

dubhthach

Wibbs wrote: »

As always Dubhthach when you post you bring real value and fantastic scholarship and on top of that make it all the clearer for the rest of us. Always a true treat to read.

Thanks for the kinds words, but I should point out I'm not a scholar, just someone who has an overactive hobby

As a note it would seem that the current "O'Brien" (aka. Baron Inchiquin) has recently done NGS (next generation sequencing) of his Y-Chromosome this confirms the link of the "Irish Type III" cluster and the marker L226 (R1b-L226) to historic Dál gCais. Baron Inchiquin has a verified lineage back at least 34 generations (32 to Brian Boru).