R-U106

Recent Burial Discoveries

Notes taken from

lain via groups as he reported on "r1b-u106@groups"

for September 2021

Please click Under-Lined item to select:

How significant the following burial is for R-U106 as a whole.


In short, there is an R-U106 ancient DNA sample from a Corded Ware Culture (page down maybe a little and click on middle of picture) burial in Plotiště nad Labem, which is about 60 miles east of Prague, just opposite the tattoo shop. The burial was dug up in the 1960s and has been dated to between 2914 BC and 2879 BC, and is of an adult male, aged 25-30. Consequently, we are looking at the foundation of R-U106 being pushed back to before 2900 BC. This is unlikely to be the R-U106 common ancestor himself, and probably not even someone who knew him, but we are clearly looking at one of his close descendants. The TMRCA for R-U106 can't be pushed more than a few centuries before this.

R-U106 burial information


Simultaneously, there is an R-L151 ancient DNA sample from another early Corded Ware Culture burial at Obříství (behind the toy shop), which is just north of Prague. First published in 2011, it is newly dated between 2911 BC and 2875 BC, and is of an adult male, aged 35-50.


Both Plotiště nad Labem (PNL001 and Obříství (OBR003) are right-sided crouched burials with their heads towards the north-west. Their only other R-U106 result is our man in Prague-Jinotice from the Unetice culture. They don't appear to have attempted typing below R-U106 for either burial. Another sample (Konobrže_26A/91, ~2900-2500 BC) is R-L151xP312, but without coverage for R-U106. Other early R-L151 burials include:

VLI011 (2881-2669 BC) xU106, xP312

VLI015 (2881-2669 BC) xU106, xP312

KO1002 (2835-2485 BC) xU106, xP312

STD002 (2882-2673 BC) xP312, xU106

VLI085 (2862-2576 BC) xU106, no coverage at P312

VLI092 (2882-2669 BC) xP312, xU106

Consequently, we are looking at a sizeable, established population of R-L151 in Bohemia during the early part of the Corded Ware Culture probably well before 2700 BC. R-L151 probably predates R-U106 by between 50 and 200 years.


So we are probably talking about ~3000 BC for R-U106 and ~3100 BC for R-L151. There's the possibility to stretch these dates perhaps 2-3 centuries further back in time without too much problem, but beyond that you start to stretch the limits of TMRCAs from modern Y-DNA and you would start to expect both more downstream clades identified in (post-)CWC remains and more basal R-L151/R-U106 clades found in modern populations further east.




Now comes the opinion part. These two results are earlier than just about anyone expected any R-L151 and indeed R-U106 results. They impart big changes to the origin story of R-L151.


Previously, the earliest attested R-L151 burial was circa 2550 BC from the upper Danube (R-U152, Bell Beaker), and the previous R-U106 burials were either our Prazan Unetice fellow, RISE98, or Lille Beddinge in the southern tip of Sweden - both from some time near 2200 BC. This pushes our R-U106 ancestry back most of a millennium to close to its foundation, meaning we can rule out any ideas of R-L151 and R-U106 not taking part in the first stages of the Corded Ware Culture. While R-L151 is later mostly found in western Europe (regions occupied by the Bell Beaker Culture [R-P312] and Single Grave Culture [R-U106]), these dates now places the foundation of R-L151 right at the start of the CWC, and significantly further east.


Another alternative hypothesis - that R-L151 and R-L52 were in western Europe before the Corded Ware Culture but had yet to be sampled - doesn't seem supported by the absence of R-L52 among pre-CWC European Y-DNA, which is now substantial, or by the autosomal DNA of these newly published burials. These new R-L151 CWC burials from Bohemia have a high fraction of autosomal DNA from the Eurasian steppe. This fraction is consistently high in the early-CWC men, and high in 10 out of the 14 early-CWC women. Hence, we can tie the first presence of R-L151 in Europe to the influx of a male-dominated DNA of steppe origin.


Hence, I suspect we can date and place the initial R-U106 starburst somewhere close to the Czech Republic. It has to be slightly older than 2900 BC, by implication it potentially began even further east. That then implies that R-L151 and R-L52 most likely formed in the "old" country to the east, but the absence of any basal clades of R-L151 or R-L52 in eastern Europe or western Asia means that they can't have been there long or built up a large population before their emigration. At the time, R-L52 was a small family branch, rather than an overbearing empire like the R-Z2105-dominated Yamnaya culture. Our ancestors were probably one of the hangers-on, and perhaps from a culture that hasn't been well-sampled yet.


The placement and timings also (weakly) suggest to me a migration over the north of the Carpathian mountains, rather than south up the Danube, as other have posited (Eupedia is a high-profile example). This is a point I've never been quite sure on, but the north migration has always made more sense to me, and I think this adds to that evidence.




So where does that leave us? This doesn't change the overall picture, namely that our ancestors migrated from the steppelands of easternmost Europe (the former USSR countries) to Europe in the Corded Ware Culture, in a wave of migration that began shortly before 2900 BC and reached the Atlantic shortly after 2700 BC. It doesn't change the fact that our ancestors then stayed in central Europe or migrated to Scandinavia, while our R-P312 brethren formed the Bell Beaker reflux and expanded to the British Isles, Iberia, Bavaria and the Italian peninsula. What it does change are the relation of the individual haplogroups to that chronology.


This is my take on the chronology. A lot of it is interpretive, so it doesn't come with much scientific rigour. A lot of it is gut feeling and subject to moderate change, even as we unravel more about the implications of these burials. The broad dates probably lie very close to the central estimates from my 2016 work:

http://www.jb.man.ac.uk/~mcdonald/genetics/table.html


The R-M269 to R-L151 chronology is still nebulous. R-L151 now probably formed between about 3300 BC and 3000 BC, and I suspect somewhere around 3100 BC, probably somewhere in the broad steppelands of easternmost Europe. Very speculatively, we might be talking about one family within a village, or a clan forming a single village, that might have been part of a wider sub-culture with loose association with the Sredny Stog culture (the culture immediately predating the Yamnaya). For whatever reason, this family or clan decided to move westwards. We don't know that reason, but it has been previously linked to the spread of the plague (Y. pestis) to Europe - our ancestors might have been fleeing the instability it caused, or were taking advantage of its effects in Europe, or both. However, our family grew rapidly from this point and for the next several centuries.


R-U106 probably formed some time between about 3200 BC and 2950 BC. I suspect somewhere close to 3050 BC. Depending on the exact time, it may already have been present in the initial migrant population, or our R-U106 ancestor may have been born during this migration period. Either way, it formed before the Corded Ware A horizon swept much of Europe, so it was early on in the migration.


I suspect that the first few generations of R-L151 remained tied together at the start, but that they parted ways somewhere very close to the Czech Republic, around 2900 BC. The first indication of geographical differences we see in R-U106 is R-Z156, and we have a descendant R-Z156 as part of the Unetice Culture appearing later in that millennium in Prague itself. So I suspect the nascent R-Z156 and the ancestors of R-S1688 (and its downstream R-U198) were left to their own devices in what is now the Czech Republic, while most of the rest of R-L151 kept pushing west. This is very approximate, because we don't sample many of the surrounding nations well.


In that westward expansion, I suspect that the ancestors of R-Z18 took the most northerly passage, ending up in the Battle Axe culture of southern Scandinavia. R-P312 probably took a more southernly passage to the Rhine Valley, and from there to the Atlantic and north-western Mediterranean coasts. Most of the rest of R-L48, many of the minor R-U106 clades, and probably our smaller brother R-S1194, probably ended up somewhere around the Single Grave culture of north-west continental Europe, especially in the regions around northern and central Germany. However, they seem to have stayed east of the P312-dominated Rhine valley until the demise of the Bell Beaker culture (~1800 BC). This R-P312 versus R-U106xP312 boundary therefore largely reflects the geographic split between the Nordic and Atlantic Bronze Ages, with a more complicated situation occurring inland.


The effects of this earlier chronology on downstream clades need some further thought. The influence here is likely to be less, as the dates don't become much more precise, and don't change as much. However, the metallurgical links supplanted from the late Unetice Culture into the early Nordic Bronze Age in the period leading up to 1700 BC probably deserve some extra attention. Clades like R-L47 could become important in this role if the timings can be made to work. Equally, this could be a source of the sporadic R-Z156 results we see scattered throughout Scandinavia that I've never got my head quite around. So there is a lot to think on.


Today's results have shown how fragile this kind of interpretation can be. Even a little data can significantly alter these chronologies. So the above shouldn't be taken as gospel - merely an interpretive view of how things stand; the most likely possibility, if you will, which could be different again tomorrow as new data comes forward, as new TMRCA calculations are made, or as we (and I in particular) learn more about the archaeological links involved.

More Thinking

Is it still reasonable to expect there were likely only one or two generations between U106>Z2265, Z2265>BY30097 and BY30097>Z381?


Yes. We have only one SNP mutation for each clade between the MRCA for R-L151, U106, Z2265, BY30097, Z381 and Z301. We don't clearly have any corresponding STR mutations, except for DYS492=12->13 between U106 and Z2265, though it remains possible that there are hidden STR mutations in the fast-mutating markers that we're not seeing because of noise created by numerous later mutations.


We expect one SNP mutation every ~83 years in BigY-700, but the rate might be slightly faster in these upper echelons of the tree where many tests are combined and the effective coverage and depth is slightly increased. We expect one Y-111 STR mutation every ~120 years (we don't have accurate rates for the Y-700 set, but mutations between these upper U106 clades would be informative).


Consequently, we expect one SNP mutation and no STR mutations to represent about 50 years on average. It must be at least one generation (circa 35 years), and it's unlikely to be more than 250 years in any single case. Because these are the larger, more populous clades, which arose during a population expansion, the times between them are likely to be more frequently towards the lower end of average.


Hence, the approximate chronology would be:

R-L151 circa 3100 BC

R-U106 circa 3050 BC

R-Z2265 circa 2950 BC

R-BY30097 circa 2900 BC

R-Z381 circa 2850 BC

R-Z301 circa 2800 BC

R-L48 circa 2650 BC

R-Z156 circa 2700 BC

All other clades, apart from R-A2150, R-S10807 and R-A8512, would form after 2650 BC. In this chronology, R-A2150 probably arose around (very roughly) 2900 BC, and R-S10807 and R-FGC8512 around 2700 BC, but it depends greatly on the number of STR mutations, which I haven't been able to easily ascertain. The whole sequence can shift forwards or backwards by about 150 years; and the distance between R-U106 at the top and R-L48 and R-Z156 at the bottom can be compressed by perhaps 100 years or expanded by perhaps 200 years. It can't expand much further because it is constrained by the 16 SNPs that should occur between R-Z156 and the R-S1894 burial at Prague-Jinotice (note this is only 7 SNPs if we discount the single reads at S1894 and S1911, and place him at R-Z304).


It doesn't matter so much for these rapidly branching clades like R-U106, but I should point out that the common ancestor for the recent R-L151 burials found in Bohemia may be more ancient than the most-recent common ancestor of all living R-L151 men. These burials may represent other R-L151 sub-clades that have died out. The short distance between R-U106 and R-L151 probably means our new burial (PNL001) lived at the same time as existing R-U106 clades. He's even likely to have known personally at least some of our ancestors, even if he isn't necessarily one himself.


I am trying to understand some of the wider implications of what the study and paper yields. There was a wide variety of haplogroups in Bohemia at that time. How and why U106 became to be associated with German speaking areas of Europe subsequently?


This is complicated, and has more to do with chance overlap with the Germanic expansions that happened three millennia later than anything else. In short, R-U106 seems to have spread over north-central Europe and southern Scandinavia. With few exceptions for minor migrations to France, eastern Europe, and probably a few other places, R-U106 seems to have remained in these confines until the Celtic era.


At this time, during the rise of Rome, our ancestors were mostly a mix of Germanic and Celtic cultures, though these are very loose terms that greatly simplify the complexity of north-west European cultures at the time. However, the Germanic expansion largely took over the Celtic lands in Europe where R-U106 is most common, so our Celtic ancestors came to speak Germanic languages.


After the fall of Rome, the R-U106 migrations continued. We know that a lot of British R-U106 are from the "Anglo-Saxon" or Viking migrations, but other British R-U106 pre- and post-date this from other non-Germanic cultures. However, all of the British Isles now speaks a Germanic language (even though we try to keep our Celtic languages alive).


So, while R-U106 was probably there at the foundation of the Germanic peoples, the association of R-U106 with Germanic language countries is more told by the story of the people who these ancestors took over, either literally or linguistically.


> Also DF96, some 1000 years younger than U106, is also said to have originated in Bohemia. So does this mean that many descendants of the original U106 man remain in Bohemia for some considerable period of time?


Perhaps. That's the most obvious explanation: that there is a genetic continuity from the Corded Ware Culture to the Unetice culture that came from it.


I will stress that we don't have direct information on R-DF96, just either its parent R-Z304 or its cousin R-DF98>S1911>S1894 (depending on how much faith you want to put in individual calls in ancient DNA). However, its spatial similarity and genetic closeness to R-DF96 means that R-DF96 probably shares the same background story.


Equally, we can't preclude that wanderlust took our R-Z304 line from somewhere near Prague in 2900 BC across the wide world and back to somewhere near Prague by circa 2000 BC, but it seems unlikely that they would be pulled back to the same point of origin.


> Do the findings of this study alter our understanding of the timing, direction and extent of the migratory movement of cultures/tribes/clans northwards and westwards of Bohemia?


Yes,


More Thinking

We can now trace our ancestral story through two significant sets of ancient DNA remains. One is the Corded Ware Culture burial published this week, which ties our R-U106 ancestry around 2900 BC close to Bohemia. The other is a Unetice Culture burial published a few years ago, which ties our R-Z304 ancestry to the same region around 2300-2000 BC. This second burial may also be S1911+ and S1894+, but this is based on very poor-quality data (only a single read of each SNP), so it could be prone to error.


So we are looking at a rough chronology of:

R-L151 circa 3100 BC, pre-Corded Ware Culture, possibly Eurasian steppe or just west

R-U106 circa 3050 BC, early Corded Ware Culture, possibly Eurasian steppe or further west

R-Z2265 circa 2950 BC, early Corded Ware Culture, probably eastern Europe, possibly still on the steppe

R-BY30097 circa 2900 BC, early Corded Ware Culture, probably eastern Europe, possibly still on the steppe

R-Z381 circa 2850 BC, early Corded Ware Culture, probably eastern or central Europe

R-Z156 circa 2700 BC, early Corded Ware Culture, probably Bohemia or nearby

R-Z306 circa 2500 BC, Corded Ware or pre-Unetice Culture, probably Bohemia or nearby

R-Z304 circa 2450 BC, Corded Ware or pre-Unetice Culture, probably Bohemia or nearby

R-DF98 circa 2350 BC, pre- or early Unetice Culture, probably Bohemia or nearby

R-S1911 circa 2150 BC, Unetice Culture, probably Bohemia or nearby

R-S1894 circa 2000 BC, Unetice Culture, probably Bohemia or nearby

R-FT18496 circa 1800 BC, Unetice Culture, probably Bohemia or nearby


The "circa" in this case could be a few centuries either way, which accounts for the lack of precise cultural assignment and location. We used to call R-DF98 the "Kings' Cluster" because we know that many royal families of Europe (the House of Wettin and apparently also the House of Bourbon) are R-DF98.


When the Unetice culture dispersed around 1700 BC, many of its descendants migrated west into Germany, where some eventually became part of the Celtic cultures that sprang up there.


More Thinking

The scientists need to find a large number of U106 and P312 in the gound in the Steppe to support a migration from the Steppe. They cannot even find one sample. And the modern dna does not support a P312 or U106 origin in the Steppe because we do not have large clusters of the earliest descendants of P312 and U106 therein. It is possible that just one L151 man arrived from the Steppe and we are all his descendants but that will be really hard to prove.


Ancient DNA samples are only found if the burials remain intact, if the soil conditions are suitable for the presence of ancient DNA, if the burials have been found, and if anyone has sequenced them... and a haplogroup only appears in those ancient DNA samples if it is a large part of the population. This means we are blind to many regions (e.g. Belarus), many cultures (e.g. cremations) and are biased towards seeing only the elite burials.


Around 3000 BC, there were probably 2-7 million people living in continental Europe, with an average life expectance of about 31 years. So over the 3300-2800 BC period of interest, there were something like 15-60 million men and boys who lived through their lives. We have ancient DNA of only a few dozen, and we are representing the entire ancient DNA content of individual countries with one or two samples... or sometimes none. There could be a million R-L151 men in the steppe before the migration, and we wouldn't see them in the ancient DNA. (We know there weren't, because we'd expect this basal R-L151 population to still be visible in modern steppe Y-DNA samples.)


Equally, we could have a million R-L151 men in Europe by 2850 BC, but we only have the few Bohemian samples to trace them, and we didn't have those last week. (This is equally unlikely, because of the time it takes to produce a million people and the haplotree structure.)


The migration of R-L151 must have happened shortly before or shortly after R-L151 formed - this is attested to precisely because we don't have basal R-L151 clades in the steppelands in either ancient or modern samples. That means that R-L151 as we know it now was only at most one extended family by the time this migration started. The chances of finding a surviving individual burial from this extended family is incredibly small, so we don't expect any U106+ or P312+ burials from the steppe.


>The CWC did not have not 100% Steppe dna. 98% of my autosomal dna is Irish. That does not mean that my S5556 branch is of Irish origin.


The CWC were a migratory group that married into the local population as they went. The burials we see - even the oldest Bohemian ones - are n-th generation immigrants, where n could be anywhere from one to more than ten. If your culture marries into a local population, you do not retain anything like 100% of your original DNA.


There's also the question of what "steppe" DNA actually is, in this case. The reference samples here are usually the Yamnaya, a R-Z2105-dominated culture that really rose to power in line with the CWC, sampled mostly from the confluence of the Samara and Volga rivers. We don't expect our R-L151 ancestors to come from this culture or their direct ancestors. That's clear from the different Y-DNA haplogroups. We just expect them to be associated.


Expecting our CWC ancestors' DNA to match exactly match the Yamanya would be like expecting your Irish autosomal DNA to match my mongrel Great British DNA exactly. Of course it doesn't, but (if we didn't know your ancestry was Irish) there are enough similarities to place your ancestry close to the British Isles. If we found someone with your autosomal DNA in Poland, we'd know it wasn't native. Similarly, we know that the R-L151 CWC burials' male ancestry wasn't from the native western or central European - it has greater affinity for cultures further east. We don't know exactly where to the east, but we know they came from somewhere around the steppelands.


> And to Keven, it is really hard to know what went on with U106 during the early years. The scientists are having trouble finding U106 samples. Perhaps U106 did not expand until 1,000 years after P312 or they were just commoners. The scientists are testing remains fron the graves of the elite.


We can tell that R-U106 expanded at roughly the same time as R-P312 from the haplotree. There is only one SNP separating R-U106 from R-L151 and only two separating R-P312 from R-L151. This means both these haplogroups split within a few generations of the main R-L151 split, and their diverse haplotrees mean they continued to split rapidly after that. We are looking at a major expansion in the size of this family that occurred over the space of a few centuries.


TMRCA estimates place that expansion within a few centuries of 3000 BC, and the ancient DNA now constrains the formation of U106 before 2900 BC. The elite burials we find from Bohemia show that there were at least some R-U106 among the elite by 2900 BC. These limits make the chronology really tight: the initial spread of R-U106 must be closely associated with the leadership of the initial phases of the CWC expansion.

It would seems to me then, having regard to your response to the first question, that a similiar study as that undertaken for Bohemia, for north-central Europe and southern Scandinavia for the period 4000 ybp to 2000 ybp would bring greater clarity to our understanding of the wanderings - albeit probably purposeful- of our R-U106 ancestors!


Yes, and various studies here are underway and starting to produce results. Sadly, very few though. Part of the problem is that cremation burials were common during this period, which means that many of our R-U106 ancestors have been reduced to piles of ashes.


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