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Sunday, September 6, 2020

Low prevalence of lactase persistence in Bronze Age Europe (Burger et al. 2020)


Over at Current Biology at this LINK. Unfortunately, this is the long-awaited Tollense Valley battle paper. Despite the obvious presence of some very interesting genetic substructures among the Tollense Valley warriors (see here), the authors have the audacity to claim that these individuals represent a "single unstructured Central/Northern European population".

One of the warriors, labeled WEZ56, belongs to Y-haplogroup R1a and shows an exceedingly Balto-Slavic-like genome-wide genetic structure. But none of this is even mentioned in passing in the paper. Indeed, according to Burger at al., WEZ56 is best classified as belonging to R1, even though the R1a classification is quite secure based on the raw data that the authors posted online.

Be extremely wary of what you read in this paper, and anything else that these scientists have published in the past and will publish in the future. Below is the paper summary:

Lactase persistence (LP), the continued expression of lactase into adulthood, is the most strongly selected single gene trait over the last 10,000 years in multiple human populations. It has been posited that the primary allele causing LP among Eurasians, rs4988235-A [1], only rose to appreciable frequencies during the Bronze and Iron Ages [2, 3], long after humans started consuming milk from domesticated animals. This rapid rise has been attributed to an influx of people from the Pontic-Caspian steppe that began around 5,000 years ago [4, 5]. We investigate the spatiotemporal spread of LP through an analysis of 14 warriors from the Tollense Bronze Age battlefield in northern Germany (∼3,200 before present, BP), the oldest large-scale conflict site north of the Alps. Genetic data indicate that these individuals represent a single unstructured Central/Northern European population. We complemented these data with genotypes of 18 individuals from the Bronze Age site Mokrin in Serbia (∼4,100 to ∼3,700 BP) and 37 individuals from Eastern Europe and the Pontic-Caspian Steppe region, predating both Bronze Age sites (∼5,980 to ∼3,980 BP). We infer low LP in all three regions, i.e., in northern Germany and South-eastern and Eastern Europe, suggesting that the surge of rs4988235 in Central and Northern Europe was unlikely caused by Steppe expansions. We estimate a selection coefficient of 0.06 and conclude that the selection was ongoing in various parts of Europe over the last 3,000 years.

Burger et al., Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years, Current Biology, Available online 3 September 2020, https://doi.org/10.1016/j.cub.2020.08.033

See also...

Warriors from at least two different populations fought in the Tollense Valley battle

98 comments:

Davidski said...

These clowns actually put Mordvins in the Slavic cluster.

Of course, Mordvins are Volga Finns.

https://en.wikipedia.org/wiki/Volga_Finns

Vladimir said...

The haplogroup 37 individuals from Eastern Europe and the Pontic-Caspian Steppe region, predating both Bronze Age sites (∼5,980 to ∼3,980 BP) not been determined?

Archi said...

They have no evidence that this is a single population, they simply did not bring them. Their PCA ranged to half of Europe https://i.ibb.co/5cNh34g/Tollense-Mokrin-PCA.png. These people, on average, have a strong shift towards WHG, and archeology finds strong ties with the Unetice culture, which I have long regarded as not entirely Indo-European (I2). The only R1a1a has a lactose tolerance gene.

The Maros culture was never associated with steppe expansion; it is a culture of purely Balkan origin. In Mokrin, they generally buried in pythos. This culture disappeared without a trace. There are R1b-Z2103 which descend from the previous Yamnayans, all the rest have nothing to do with the steppe, these are typical Balkan(I2a)-Anatolian(J2b) subclades. Naturally, they are not Indo-Europeans.

Hence it follows that the conclusions that the mutation of lactose tolerance is not from the steppe and its prevalence in the Bronze Age are premature.

Rob said...

Maybe it’s the preamble before a new paper will then focus on the main population genetics and and structuring of late Bronze Age northern Europe?

Archi said...

A characteristic feature of Tollense is that it does not contain Eastern European mtDNA, except for two cases (WEZ59 XY U5a2b1a R1b1a2a1a2, WEZ61 XX U4b1b1). That is, on the female line, there are also all from TRB/Globular Amphora and Unetiсe.

Rob said...

@ Archie

“ This culture disappeared without a trace. There are R1b-Z2103 which descend from the previous Yamnayans, all the rest have nothing to do with the steppe, these are typical Balkan(I2a)-Anatolian(J2b) subclades.
Naturally, they are not Indo-Europeans.”

Z2103 is the most likely vector of post-Anatolian IE in southeastern Europe
Given that I2a1, J2b and R1b-Z2103 are all found in the Balkans to this day; your statement is obviously false

zardos said...

They want to avoid, at all costs, that there might be an impression of "ethnic warfare", which is of course not that easy to deduce with certainty anyway, since various alliances are thinkable. Yet the overall impression is still rather like that, looking at the remains.
Another issue is how you define "Northern-Central European", because the term can be used very inclusively. I mean even today people might have a wider or narrower definition of what "Central European" ancestry is about, like it could include anything ranging from Northern German-Dutch, over Belgian-French, to Swiss Italian to Hungarians-Czechs, probably even Poles-Slovaks. If you use such a wide definition of Central European, and they even say North-Central European, like including Scandinavians too, they are surely right, but that's beside the point.
The third is that they mainly relied on strictly statistical methods, without using common sense and their senses. If they processed the data right, I wonder what magnitude of differences these insufficient methods need to come to the conclusion of significant grouped variation.

Another problem are the referene samples in their tree (Fig 2) which is very, very poor for a recent study. I mean CEU/Utah whites as the main related sample! No modern Germans (not dreaming of from different regional German samples), Poles, Czechs etc. That's ridiculous!
It just shows how important modern regional samples are for a proper comparison.

Concerning LP, its now quite obvious that we deal with an exremely intensive, hard selection for it in quite recent times. The main explanation for this is, considering that the LI individuals did consume milk and milk products for a long time before successfully, that for a specific phase large portions of the European population must have primarily relied on milk consumption for survival.

zardos said...

Looking at both Tollense and Mokrin, its interesting to add that both populations were so far dominated by specific clades of R1b and I2a. Looking at the later and modern populations in these regions, this looks not like continuity at all, but rather like a major upheaval coming after these populations existed.

Vinitharya said...

Yeah, I got a strong impression of fishiness with the haplogroup assignments; not only are there no proto-Baltic or pre-Slavic haplogroups, there are no Germanic clades either, besides the relict M223 that was assimilated by the early Germanic tribes. No I1, no U106, no R1a of any kind. This is more like 'band of Celtic adventurers far from their Rhenish home versus a bunch of Funnelbeaker remnants who have managed to survive assimilation for over a millennia', as an American with low tolerance for cultural cringe, I am disgusted that such thinking has crept into science. I really have little to say about the lactase persistence, except for the fact that lactose tolerance strongly correlates with higher Steppe influence among European populations.

Rob said...

@ Zardos

''Looking at both Tollense and Mokrin, its interesting to add that both populations were so far dominated by specific clades of R1b and I2a. Looking at the later and modern populations in these regions, this looks not like continuity at all, but rather like a major upheaval coming after these populations existed.''

What are you talking about ?
Instead of blind speculations, read my comment - ''I2a1, J2b and R1b-Z2103 are all found in the Balkans to this day; ''

As for northern Germany- what LBA/ Iron Age data are you referring to to make your deduction ?

zardos said...

@Rob: Yes, they are there, and were even stronger before the Germanic and Slavic expansion, but the later Roman samples are still quite different overall. And I think the difference would be even bigger, if there wouldn't have been related people coming in.

Concerning Northern Germany, I just have to refer to what Vinitharya posted. Together with the unusually high WHG-percentage, this can only create an impression of a population which was, to a large degree, replaced in prehistorical times.

ambron said...

Archi, it is interesting that in the Viking study, lactose tolerance was also associated with the flow of Balto-Slavic genes.

zardos said...

@Rob, No 2: The exact subclades of I2 from Mokrin are particularly noteworthy, not really common in modern Pannonian-Carpathian-Balkan populations at all.

AWood said...

There's no agenda here. The most plausible answer is this is an army from the south who was securing a trade route, they may not have been local at all. Southern Germany seems the most plausible to me based on the R1b haplogroups. I can't really speak for the I2, since M223 is found everywhere in Europe, but at very low frequency today. Has anyone taken a look at which ones are southern and which ones are local? Is there a pattern? Also, keep in mind 170 different skeletons were determined, out of a battle of several thousands. We are just looking at 17 (or so). No reason to think I1 or more R1a couldn't pop up. However, if the implication is that this is a southern army coming to secure trade against a resilient group of locals, there is no guarantee, especially if these are remnant I2-M223 TRB who held out thousands of years.

AWood said...

Actually northern Europe has plenty of R1b. Perhaps not Baltic Europe, but Scandinavia, in particular southern Scandinavia and northern Germany has a lot. What was thought in 2005 was diverse haplotypes of R1b meant age. What this really meant was diverse branches of R1b (ie: L21, U152, U106, DF27..etc) which was the result of layered migration/movement of people. The less diverse branches such as R1a, indicate the bottleneck.

Matt said...

I'd suggest that if they wanted to find substructure among European populations using formal stats, they perhaps ought to have used: f4(TestTollense1,Ust_Ishim;TestTollense2/TestModern,Mbuti).

f4(TollenseTest1,TollenseTest2;Test,Yoruba) could become dominated by differences in Basal Eurasian between Test 1 and 2.

I find that rough equivalents made from f3 stats of f4(Modern1,Ust_Ishim;OtherModerns,Mbuti) tend to reproduce the relationships from Fst scores fairly well (without much effect of strong local drift), because de-emphasizes the Basal Eurasian portion.

I would guess that if you are looking at the later affinities, you might want to deemphasize the effect of the Basal Eurasian portion dominating the signal.

(Or the effect whatever the Basal Eurasian type pattern really is, geneflow with unsampled African population or something, if it never actually existed.)

Then also compare to (TestModern1,Ust_Ishim;TestTollense/TestModern2,Mbuti) to check significance against known present day structure, rather than just against Z=3. It would probably be useful to check them against the size of effects in present day known structured populations as well, rather than just use an arbitrary significance threshold? We might be interested in if the Tollense samples are structured *relative* to modern day structure rather than if they are structured relative to significance tests?

As to whether there was ever, even since the Middle Neolithic at 3000, really a single unstructured North/Central European population, it seems hard to say.

Open Genomes said...

WEZ56 is actually Z662;CTS11197;PF6225+
C>A (1A)
And therefore "Norse-Balto-Slavic" R-Z283.

Samuel Andrews said...

It will very interesting to see more Tollene DNA samples. Amatuers here will be able to figure out exactly what they are. Was this battle madeup of soldiers from many tribes, maybe some from one confederation (Y DNA I2a), where did they all come from. Stuff like that could all be answered which would help understand Tollense valley Battle.

From small samples we have we already know one soldier is from Spain, one from France, one from Northeast possibly Belarus. And that the rest come from an extinct regional population who carried mostly Y DNA I2a2 but could have been represented by many tribes.

Samuel Andrews said...

They conclude Lactose persistence can't be from Pontic Caspien Steppe because no ancient samples from there have it. They don't understand the concept of natural selection. Most selected traits were at once at ~0% in the population they originated in.

Lactose Persistence probably originated in the Steppe ancestor of Corded Ware, began to be selected for in Corded Ware period, was selected for much more heavily from 1500-500 BC.

Archi said...

@Rob

"@ Archie @ Zardos "

Learn to read. I wrote the truth, you wrote only your fantasy, and you also pass your own fantasy as truth, and you attribute a lie to me. The lie is that Maros culture has continued on. It is a lie that Maros/I2a/J2b are of Indo-European origin. Maros culture was pushed here from the Balkans under the pressure of Vatya culture. Naturally, after the Maros culture, enormous genetic changes took place here.

See Yamnaya/Z2103 in previous time:
Bronze Vucedol Croatia Beli Manastir-Popova zemlja [I3499 / GEN72] 2884-2666 calBCE (4176±28 BP, BRAMS-1304) M R1b1a1a2a2 [CTS1078 / Z2103] T2e

Bronze Yamnaya Bulgaria Benkovski [BEN3] 5000-4500 y.a. H
Bronze Yamnaya Bulgaria Golyamata Mogila, Popovo [POP1] 5000-4500 y.a. T2a1b1a
Bronze Yamnaya Bulgaria Golyamata Mogila, Popovo [POP3] 5000-4500 y.a. U2e1a
Bronze Yamnaya Bulgaria Golyamata Mogila, Popovo [POP4] 5000-4500 y.a. U5a1
Bronze Yamnaya Bulgaria Riltsi [RIL3] 5000-4500 y.a. K
Bronze Yamnaya Bulgaria Ovchartsi [OVI2] 5000-4500 y.a. K
Bronze Yamnaya Bulgaria Ovchartsi [OVI3] 5000-4500 y.a. U/K

Samuel Andrews said...

Lactose Persistence is from Indo European languages.

It comes from Corded Ware who was the main spreader of IE languages. Or more specifically, it began to be selected in Corded Ware.

The first consistent instance of Lactose Persistence are in Corded Ware, Bell BEaker, Andronovo.

Then, it was selected for in Mid-Late Bronze age, in former Corded Ware & Bell Beaker territory, as well as in some dairy dependent populations in India.

@Vinitharya,
"I really have little to say about the lactase persistence, except for the fact that lactose tolerance strongly correlates with higher Steppe influence among European populations."

Samuel Andrews said...

The Bronze age Serbian samples are interesting similar to modern South Slavs. They will probably overlap with modern West Balkans in West Eurasia PCA.

Even though, this isn't population continuity from 2000 BC to present day. Iron age Balkans were probably significantly less Steppe than these Bronze age samples. Or maybe, in Roman era there was Greek admixture in West balkans which pushed modern Slavs south of Bronze age Balkans-Slavic continum.

Samuel Andrews said...

The none-Lombard samples in Hungary, cluster close with Bronze age Balkans but also have Near Eastern admixture which can be linked to admixture in Roman era.

This suggests a norm was established in West Balkans in 2000 BC which continued till Roman era when minor Near East admix came in.

But, it's very likely the "East Med" shift in modern West Balkans comes mostly from Greece/East Balkans considering there's E-V13 there today.

Modern South Slavs, aren't perfect Bronze age Balkans-Slavic mix, there's "East Med" admix either from Greece or Near East.

Michales said...

Accidentally I found PDF with full article:)

https://drive.google.com/file/d/1rvi8pZJJabzFdysdR2Oo3kx4Rf8o59yV/view?usp=sharing

Ioannis Gavras said...

@ Samuel

East Med shift in balkans exist even during CA/BA period.West Asian like admixture(CHG/IRAN N) and i am pretty sure the IA balkan period was probably mostly of EEF/Steppe + adittional CHG/IRAN N input.It dosn't necessary means it comes from Greece.It was actually Romans who dominated later these lands and getted them into Roman-Byzantium empire(Vlachs etc).The Greeks never went in balkans,it was actually vlachs and slavs who settled down to Byzantium.Not any serious reason for Byzantines to move north and mix with natives.That middle east autosomal has CA/BA/IA relation.

Davidski said...

@Michales

This is an open access paper. The PDF is freely available at Current Biology.

https://www.cell.com/current-biology/pdfExtended/S0960-9822(20)31187-8

Archi said...

So far, we can conclude that I2a were defeated in the Tollense battle.

Samuel Andrews said...

I think the older Chalcolithic Southwest Asian admix only impacted Greece and Sicily. Bronze age samples from Serbia, Croatia lack it. Early Bronze age samples from Bulgaria lack it. As does single Iron age sample from Bulgaria. The "East Med" shift in modern Balkans should be from Roman period.

Ioannis Gavras said...

https://docs.google.com/spreadsheets/d/1a5byEejcptHST6X2L6Exb60uGXnyI6tZYI7HWG-EkkE/edit#gid=0


The balkan samples from Bulgaria are kinda mixed.Look their farmer ancestry.They are not just barcin N but they got tepecik çiftlik N admixture.The later farmer site inclundes components like Iran/CHG and Levant N witch shifting them more east.These components might be in Bulgaria since the Neolithic.

zardos said...

@Sam: It doesn't really matter which group had a high proportion of LP originally, but by all we know, it seems it wasn't the steppe people. This is even more clear than for the depigmentation of eyes and hair, by now its practically out of question.
But on the long run, selection favoured the LP trait in steppe derived populations, which were already steppe-neolithic mixed. Like you said yourself, the important aspect was the selection. Question is which changes in lifestyle led to the enormous acceleration of the selection process. In the paper they name some aspects of this, and argue for a combination, which I think is basically correct. But in the end, it all can only lead to the conclusion that for a specific time span the dependency on milk and especially fresh milk, must have been, in some people of the North, very extreme.
And most of the strongest selection happened not before the Middle Bronze Age or in so far unsampled populations - I rather think it was a change of lifestyle that spread together with cultural innovations, which led to a shift in the selective regime.

@AWood: This was too big, it was an invasion or at least an attempt to conquest, which might have failed. Looks like the local tribals used simpler weapons, especially wooden weapons and arrowheads of stone en masse to lure the incoming warriors from the relative South in a trap, where they butchered them. The failed conquerors were better equipped and had more elobarated gear. Like much more Bronze weapons and tools, horses and armour. They lost primarily because they were in an ideal position for being shot by volleys of arrows, in a bad defensive position after the surprise attack by the locals.

Ioannis Gavras said...

There is also a post-Mycenaean West Asian migration to the Balkans.This east med shift is not just from Tepecik farmers.Thought the Slavic mix has reversed effects of it to some extent.I am sure more samples from balkans in the future especially from Bulgaria will show such an admixture more clearly.

Archi said...

@zardos

This is all just speculation about the weapons and causes of war, there is no data on why the alien army came here. The reasons for the war and who fought at all are unclear. Given the subsequent catastrophe of the Bronze Age it could be a war because of the Bronze shortage, or maybe the opposite, the northerners were moving to the south, maybe the cause of the war was the collapse of the Tumulus culture, maybe ...

Open Genomes said...

So far, WEZ15, the highest coverage sample, is I-Y3672*, specifically I-Y3672 Y7244- Y4332- Y10648- YP4799- Y125786- Y53080- Y13008- Y13321- Y33734-.

The tMRCA of I-Y3672 in YFull is 3400 ybp, 1400 BCE, and the radiocarbon dates for the battle are 1300-1250 calBCE. Pretty close in time.
https://yfull.com/tree/I-Y3672/

To me, it seems that this group appears to be rather anachronistically "Proto-Celtic", meaning that their descendants appear to be mostly of Celtic origin, with some exceptions. Perhaps we can say that all present-day I-Y3672 individuals are descended from this very "tribe", or at least, very close relatives of theirs.

Samuel Andrews said...

@zardos,

You're right to point out this doesn't mean the Lactose mutation originated in Indo European Steppe population. And that it rose to modern high frequencies long after the IE migrations.

But the IE migrations, specifically Corded Ware, is what spread the mutation across most of Europe.

If not for the IE migrations, the high frequencies of the mutation today wouldn't exist. So, population movements do play an important role. In that, the mutation had to be spread by a population and that population was Corded Ware.

Also, Lactose Persistant mutation began to be selected during the Corded Ware period. Because it pops up at a consistent but very low frequency in Bell Beaker and Fatyanovo.

How did Fatyanovo in Moscow and Bell beaker in Britain both get the same mutation? IE migrations.

zardos said...

@Archi: The reasons for the campaign are unknown, but it was an invasion and the course of events was suggested by archaeologists as a probable scenario. They found a lot of rather primitive stone arrowheads, even sticking in the bodies of well-equipped warriors, of which a higher proportion had state of the art bronze arrowheads, even Tüllenpfeilspitzen.

So probably it was a revenge campaign or whatever, but it seems to have been an invasion rather and the most likely winning side of local tribal warriors used simpler, more often wooden and stone weapons.

AWood said...

@zardos

I don't really follow how you arrived at this conclusion of who won and who didn't.

Rob said...
This comment has been removed by the author.
Michalis Moriopoulos said...

@Davidski

Will you be adding the samples from this paper (Tollense, Mokrin, etc.) to the G25?

Davidski said...

@Michalis

Yes, if and when the genotypes become available.

ambron said...

Archi, or maybe I2a and R1b were defeated ... After ancient battles, the victors collected and burned own fallen warriors. This one R1a drowned in the swamp and was not found.

Rob said...

@ Zardos

''
Concerning Northern Germany, I just have to refer to what Vinitharya posted. Together with the unusually high WHG-percentage, this can only create an impression of a population which was, to a large degree, replaced in prehistorical time'''

Vinitharya's deduction that ''This is more like 'band of Celtic adventurers far from their Rhenish home versus a bunch of Funnelbeaker remnants '' is difficult to sustain
First off, how do we get TRB stragglers in 1200 BC ? TRB was 'gone' by 3000 BC. The I2a haplogroup in Tollensee isn;t even from TRB, but from Wartberg

The claim that R1b are Celtic adventurers is speculative atm. We would have to ascertain with further sampling from 'normal' burials related to civilian settlements to determine what native populations looked like. But this combination actually looks unsurprising for LBA Germany


''to a large degree, replaced in prehistorical times.''

Obviously modern Germans are different. But modern Germans are actually irreevant for BA - IA Germany

The large region of mittelEuropa, from northern germany to central Balkans experienced a ping-pong of shifts every few hundred years expanding west to east or viceversa (Unrfield, Tumulus C., Halstatt, etc)
So shifting ''cultures'' and supremacy is the rule.
But how this affected genetic make up we are still in the dark. Thus, claiming a marked shift is premature. In fact, the largest change in central Europe occurred in after 400 AD Long after Tollensee battle, after which it is likely to have been followed by relative population continuity


Balkans:
''Yes, they are there, and were even stronger before the Germanic and Slavic expansion, but the later Roman samples are still quite different overall. And I think the difference would be even bigger, if there wouldn't have been related people coming in.''

Which Roman samples is that ? Lombards , which are obviously foreign.?
Again shifts are the rule in this part of Europe, but grosso modo , I am actually suprised that they are fairly similar to the pre-balto-Slavic component in northern Balkans. As you now say, a similar population lived around, withoug implying absolute continuity in a specific locale

Rob said...
This comment has been removed by the author.
Rob said...

@ Archi


'' The lie is that Maros culture has continued on.''

No shit. All cultures end or transform. the Maros culture ended ~ 1600 BC
But you said 'vanished without a trace', which is clearly not true


''It is a lie that Maros/I2a/J2b are of Indo-European origin.''

Did i say they are 'of IE origin' ?
Re-examine what you said ''Naturally, they (collectively, incl R1b-Z2103) are not Indo-Europeans.''
By 2000 BC, these people *might* have spoken some form of IE. We dont know they didn't. It might have been one of the idioms they spoke.


See Yamnaya/Z2103 in previous time:
Bronze Vucedol Croatia Beli Manastir-Popova zemlja [I3499 / GEN72] 2884-2666 calBCE (4176±28 BP, BRAMS-1304) M R1b1a1a2a2 [CTS1078 / Z2103] T2e

Bronze Yamnaya Bulgaria Benkovski [BEN3] 5000-4500 y.a. H
Bronze Yamnaya Bulgaria Golyamata Mogila, Popovo [POP1] 5000-4500 y.a. T2a1b1a
Bronze Yamnaya Bulgaria Golyamata Mogila, Popovo [POP3] 5000-4500 y.a. U2e1a
Bronze Yamnaya Bulgaria Golyamata Mogila, Popovo [POP4] 5000-4500 y.a. U5a1
Bronze Yamnaya Bulgaria Riltsi [RIL3] 5000-4500 y.a. K
Bronze Yamnaya Bulgaria Ovchartsi [OVI2] 5000-4500 y.a. K
Bronze Yamnaya Bulgaria Ovchartsi [OVI3] 5000-4500 y.a. U/K''

Relevance ?

Davidski said...

@All

Can we link these sample codes to very specific archeological descriptions of the fallen and actually figure this out?

That is, can we figure out with some certainty whether these people belonged to the same or different armies, and, in particular, is there anything specific about WEZ56 that suggests that he may have been on the opposite side of the conflict to the R1b/I2a warriors?

Copper Axe said...

When I first read about the Tollense valley I figured it was an army of Tumulus/Urnfield together with Lusatian allies (as they seemed to be very "Celtic" influenced) fighting against a less technologically advanced population from the north. Maybe a giant annual caravan route that got surprise attacked?

zardos said...

@Sam:
"Also, Lactose Persistant mutation began to be selected during the Corded Ware period. Because it pops up at a consistent but very low frequency in Bell Beaker and Fatyanovo.

How did Fatyanovo in Moscow and Bell beaker in Britain both get the same mutation? IE migrations."

Yes, but at that time most migrations were "IE migrations", so this is beside the point. It seems that some Neolithic groups had higher, but still very low frequencies of LP when the steppe people came in. And because there were many mixtures and back migrations, like Abashevo-Sintashta is an absolutely clear case of a back-migration, the allel spread and since the Northern steppe derived people adopted a specific lifestyle and more migrations followed, selection did its work.
But the allel was introduced in all likelihood by Neolithics, like depigmentation, and it spread only in a later phase, after much of Europe was already IE.

@AWood: "I don't really follow how you arrived at this conclusion of who won and who didn't."

Because we have many, many stone arrowheads around the battlefield and a lot of highest class, best equipped warriors ending up in the river with these arrowheads around them and in their body. They were first brought into disarray, then pushed together, and last they fought for their life at the shore or already in the river, where they were, quite often, punched to death by rather primitive wooden weapons. Like the bats which were found and are so far extremely rare in the archaeological record, but appear also in more modern Celtic contexts from Britain by the way.
Many of those lying on dry ground were robbed, but those under water not. And a lot of them look like better equipped, high status or professional warriors.
They were caught in a surprise attack and it rained stone arrowheads on them, after which the attackers pushed them into the water and punched them to death with bat and club like weapons for the most part, that's just what the findings tell us.
They could have robbed some from the dry land and throw them into the river afterwards - so not all which have no good tools and weapons might have gone to battle like that.

And yes, its highly likely that a large portion of the winning side's victims, of which some must have existed, considering how bloody and desperate the fight must have been, was taken away and buried properly to their customs elsewhere. But looking at how the well-equipped warriors were slaughtered at and even in the river, its at least highly likely that the arrow shooting locals with bats and clubs made the day.

@Rob: I don't meant everything he said, but the general notion of the "lack of Germanicness" which can be attributed to the findings. One haplogroup looks quite Celto-Germanic, rather Celtic by the way.
WEZ15 = (?) https://www.yfull.com/tree/I-Y3672/

How is his autosomal and archaeological profile?

Aram said...

Archi

The story of J2b2a in Europe is not solved yet. It will not change the general picture of PIE story. Nevertheless it will add some nuances if we learn that it came via Steppe or was a Cardial ware related Neolithic marker integrated into Steppe groups.
A post Neolithic migration is also possible but less likely.

zardos said...

Looks like WEZ15 is more Eastern shifted in comparison to the average. Funny considering how his paternal haplogroup being distributed nowadays.

Archi said...

@zardos

Stone arrows were used by everybody, for example, the most civilized Mycenae. Bronze was very expensive and rare, and it is absolutely unclear why bronze arrows should be used in the absence of the armours.

Rob said...
"'' The lie is that Maros culture has continued on.''
No shit. All cultures end or transform. the Maros culture ended ~ 1600 BC
But you said 'vanished without a trace', which is clearly not true"

Don't be verbiage. Maros culture was replaced by Tumulus culture, which is not related to it in any way. There is no continuity.

"Re-examine what you said ''Naturally, they (collectively, incl R1b-Z2103) are not Indo-Europeans.''"

I did not say that. "Not Indo-Europeans" origin related only to the last phrase (I2a/J2b), however, R1b-Z2103 lived so long in the environment of non-Indo-Europeans that at that time also could stop being them. But again, I did not write that it was Non-Indo-European.

Mesolithic Iran Hotu Cave [I1293 / Hotu IIIb] 9100-8600 BCE M J (xJ2a1b3, J2b2a1a1) [J2a-CTS1085]
Neolithic PPN Iran Tepe Abdul Hosein [AH2] 8205-7756 calBCE (8833В±41 BP, MAMS-25472) M J2b-M12*
Copper Iran Hajji Firuz [I4241 / F10 B1 S3] 6016-5899 calBCE (7080±30 BP, PSUAMS-2163) M J2b
Copper Iran Hajji Firuz [I4349 / F11 4 merged with F11 B3 3] 5887-5724 calBCE (6915±40 BP, PSUAMS-2126) M J2b
Copper Iran Seh Gabi [I1662 / SG7] 4831-4612 calBCE (5860±40 BP) M J (xJ1a, J2a1, J2b) [J2a2-PF5008 (xPF5058)]
Bronze Croatia Veliki Vanik [I4331 / VV1] 1631-1521 calBCE (3305±20 BP, PSUAMS-2257) M J2b2a
Bronze Early Bronze Jordan 'Ain Ghazal [I1730 / AG 84_30] 2489-2299 calBCE (3925±31 BP) M J (xJ1,J2a, J2b2a) [J2b1-PF7331]
Bronze Canaanite Lebanon Sidon [burial 63] 1600 BC M J2b

Archi said...

@Davidski

There is no description of the samples. The disgusting feature of this paper is that it does not describe the archeology of these samples, where they were found, the archeological context those samples that everyone was waiting for, because you will not get such information anywhere else as from this paper.

zardos said...

In an important article about the Tollense battle, the authors wrote that the arrowheads of bronze with a cup ("Tüllenpfeilspitzen") being very rare in the North, only 28 (!) have been found at all in the larger region of Mecklenburg-Vorpommern. So they were not unknown, but very rare. Yet in Tollense 49 were found so far, with much more being expected and parallels to regions to the South, Southern Germany, in Czechia, Poland and more Southern Eastern Germany. From Scandinavia not a single such arrowhead was ever found so far!

In the same article they argue for a trap and that the attackers on the column fired volleys of arrows from higher ground on the largely helpless invasion party, which was taken by surprise.
Many of those dying from this invading groups were high status and professional warriors, with highest quality and very expensive goods.

The article was produced for a large exhibition:
https://www.researchgate.net/publication/284142174_H_Meller_Krieg_-_eine_archaologische_Spurensuche_In_H_MellerM_Schefzik_Hrsg_Krieg_-_eine_archaologische_Spurensuche_Begleitband_zur_Sonderausstellung_im_Landesmuseum_fur_Vorgeschichte_Halle_Saale_6_No

Unfortunately they don't use examples from the record which could be connected to the genetic results. Particularly the warriors with the elaborated equipment and pieces of jewelry, like golden hair rings, would be of particular interest, because they are highly likely to represent the elite of the invading army.

Archi said...

@Zardos, should I remind you that yet Hercules was running with a wooden club? Wooden batons were quite common weapons for both Mycenae and Doriens.
For example, the basis of weapons of the Mycenae army were not even archers, but slingers, who fought in general absolutely naked.

Rob said...
This comment has been removed by the author.
Rob said...

@ Archi

'' Maros culture was replaced by Tumulus culture, which is not related to it in any way. There is no continuity.''

That region was a keleidoscope of groupings, no culture ''continues on' . Its shifts every few hundred years. But there is genetic, ergo population continuity.
Nobody is claiming there is any direct filiation, perhaps except for youreself


''Mesolithic Iran Hotu Cave [I1293 / Hotu IIIb] 9100-8600 BCE M J (xJ2a1b3, J2b2a1a1) [J2a-CTS1085]
Neolithic PPN Iran Tepe Abdul Hosein [AH2] 8205-7756 calBCE (8833В±41 BP, MAMS-25472) M J2b-M12*
Copper Iran Hajji Firuz [I4241 / F10 B1 S3] 6016-5899 calBCE (7080±30 BP, PSUAMS-2163) M J2b
Copper Iran Hajji Firuz [I4349 / F11 4 merged with F11 B3 3] 5887-5724 calBCE (6915±40 BP, PSUAMS-2126) M J2b
Copper Iran Seh Gabi [I1662 / SG7] 4831-4612 calBCE (5860±40 BP) M J (xJ1a, J2a1, J2b) [J2a2-PF5008 (xPF5058)]
Bronze Croatia Veliki Vanik [I4331 / VV1] 1631-1521 calBCE (3305±20 BP, PSUAMS-2257) M J2b2a
Bronze Early Bronze Jordan 'Ain Ghazal [I1730 / AG 84_30] 2489-2299 calBCE (3925±31 BP) M J (xJ1,J2a, J2b2a) [J2b1-PF7331]
Bronze Canaanite Lebanon Sidon [burial 63] 1600 BC M J2b'''

Yes I know that, but thanks for summarizing it for everyone

@ Zardos
i guess they look west European, autosomally & uniparentally. But i havent looked at it in detail
The curious thing is - proto-Germanic, with I1 (and possibly U106) also looks ''west European''

zardos said...

I know that and don't mind, actucally expect that the invading party had clubs and bats too, I'm also pretty sure that the locals had bronze weapons as well, however, the percentage and distribution points to a difference: The invading army was "state of the art" with a high percentage of elaborated weapons, armour, tools, jewelry etc., everything point to a more Southern origin, while the locals had more the appearance of a tribal contingent, probably local farmers and herders for the msot part. Surely under leaders, which in turn might have their band of warriors too, with armour and everything.
But some of the dead in the river look really like experienced high ranking and half-professional warriors with, I repeat it again, state of the art, best available equipment. And to me it doesn't look like both sides were like that, which would contradict the general record, but those which went into the trap were.
I doubt the local party would have went into the trap of the foreign invaders.

So everything considered, Tollense shows parallels to the battle in the Teutoburger Forest or similar scenarios, in which an otherwise superiour army was lured into a trap, caught in it and slaughtered. There was so much material that the locals couldn't carry everything away, they took only the most valuable goods from the defeated and thanks to the water, some were left untouched altogether.

Rob said...

It seems like theyre remains pulled out from a 2,5km stretch of the river. So it is interesting that the males follow such as distinctive dichotomy of R1b-L51 and I2a2a1a

So, they either represent 2 opposing sides, from both of which the dead were 'buried' in the river; or
They are both from one (?losing) side, which was disposed off into the river

When data comes out, genome-wide properties can be compared against Y-DNA, so that might help clarify

zardos said...

I think there are just two options:
- The rather South-Western shifted elite warriors were accompanied by (local?) allies which were rather more North-Eastern.
- The fight was so intense that when closing in, the fighting warriors of both sides died in the river and at its shores, due hand to hand combat.

I think both is possible and likely. I mean the whole thing is so big, why not thinking of a local tribe which called for help from reknowned allies to the South. Their opponents knew they were coming, prepared a trap and the allies went right into it. Doesn't sound that far fetched at all, considering how large the Bronze Age networks were.

I also have to say that this battle might have been important for the more Southern alliance, because I guess that such a loss of well trained and equipped warriors was hard to recover from. I guess they might have become an easy or at least easier pray for other neighbours. Because the remains found so far might represent only a very small fraction of the true losses and thinking about it, that was an enormous loss of humans and material. Hard to recover from I guess. They were not the Roman army after all...

There is a difference noted by the record, that many of the bodies which were not under water seem to have been robbed. We can't know for sure, but I think its unlikely that they would have left the own dead to rot at the place, after having robbed them. I mean we can't know for sure, that might have been their custom, but I doubt it.

The most interesting remains being found from the area which was below the water surface and therefore harder to access by the winning party.

Samuel Andrews said...

@zardos,
"Yes, but at that time most migrations were "IE migrations", so this is beside the point."

There were a set of migrations which spread IE languages across Eurasia. Those are the only IE migrations.

"It seems that some Neolithic groups had higher, but still very low frequencies of LP when the steppe people came in. "

No modern study has found LP allele in Neolithic farmers. The first consistent instances of LP come from Corded Ware and Bell Beaker.

Which links the spread (not high frequency) of LP allele to IE migrations. Because CWC and BBC were the main spreaders of IE languages across Europe.

Samuel Andrews said...

@zardos,
"But the allel was introduced in all likelihood by Neolithics, like depigmentation, and it spread only in a later phase, after much of Europe was already IE."

LP was introduced in most of Europe with IEs. It's possible it ultimately originated in a Neolithic farmer population. But, it was spread across Europe, introduced to most parts, by IEs.

There is a direct connection between LP and IEs, in that they dispersed it and made it possible for it to be selected in later time.

Ric Hern said...

Sounds a bit like Bog Bodies or some ritual related to it. If so then areas from Denmark to Netherlands, Britain and Ireland comes to mind...

Samuel Andrews said...

I see interesting theories on what Tollense battle represents.

Everyone except the experts already know....

Many Neolithic to Bronze age European societies had a single Y Chromsome.....

So the fact there's only R1b L51 and I2a2a1a in the soldiers really suggests this was battle between two different populations one R1b and and one I2a.

It would be really interesting if one population came from a far away. We already know there were mercenaries from Spain, France, and Lithuania.

Samuel Andrews said...

@zardos,

You think I2a2a1a population is primitive native population, the R1b is Bronze age invaders from Southwest?

Combining archeaology with DNA, we should be able to figure that out.

Davidski said...

@zardos

It might be useful to cross check, if possible, my autosomal analysis of some of the Tollense samples with the Y-haplogroups from this paper.

https://eurogenes.blogspot.com/2017/10/tollense-valley-bronze-age-warriors.html

WEZ15 I2a1b1a1b1a1a
WEZ24 I2a1b1a2b1a2a1a1a1a3a2~
WEZ35 R1b1a1b1a1a2
WEZ39 I2a1b1a1b1~
WEZ40 R1b1a1b1a1
WEZ48 I2a1b2a2a3
WEZ51 I2a1b1a1b1a1a~
WEZ53 R1b1a1b1a
WEZ54 R1b1a1b1a1a2
WEZ56 R1a1a1b1~
WEZ57 R1b1a1b1a
WEZ58 I2a2a
WEZ59 R1b1a1b1a1a2a1a
WEZ64 I2a1b1a1b1a
WEZ71 I2a1b1a1b1~
WEZ83 I2a1b1a1b~

And go from there...

Archi said...

@Samuel Andrews

"Because CWC and BBC were the main spreaders of IE languages across Europe."

The BBC were not the spreaders of IE languages in Europe.
They were not native speakers of Indo-European languages at all, but became so from mixing with CWC. The distribution of IE languages by R1b already took place mainly in the Iron Age.

zardos said...

@Samuel: I guess LP was present at low frequency in much of the Neolithics. So if you like the back migration of mixed steppe people brought it at higher frequency to the East probably. The main factor was a lifestyle change postdating the steppe peoples original expansion.
The situation seems to have been different in South Asia, there steppe derived people were probably the main DIRECT vector for the spread of LP from the start.

"You think I2a2a1a population is primitive native population, the R1b is Bronze age invaders from Southwest?"

That wasn't necessarily my main point at all, because I don't think that the haplogroups must be that decisive. Even more, there could be different alliances.
And I wouldn't call them primitive at all, just simpler, rather poorer and backward people in comparison to the elite warriors in particular, which had wide ranging connections and gear from some of the best smiths of their time, with artefacts which never made it to Scandinavia.

WEZ35 is R1b-P312 and most put him into a more Western grouping.

All the clearly more Southern shifted individuals are R1b (53, 54, 57). So there is a tendency for the more Western and partically Southern group to have more R1b, but its not decisive?

Probably the main local group was still I2a, with some R1b married in, with the majority of the Southern/Western shifted elite warriors coming as allies being R1b and the opponents more exclusively I2a? The sample size is too small, archaeological context lacking, but it might be that way.

Does anyone have data from find spot WEZ28? Because thats the find spot of the pourse. This article is quite interesting and repeats the story of the paper for the exhibition (popular scientific newspaper):
https://www.spektrum.de/news/auf-kriegszug-gen-norden/1680250

Romulus said...

The victors bury their dead, the losers don't. The losers were Urnfield, who we know carry I2a2 and come from an area rich in P312. Victors were Nordic Bronze age groups? Who had culturally diverged from Urnfield many centuries prior when they started making their own metal work, metal deposits potentially being what the Urnfielders were after. Archaeologists suggest that the Urnfield expansion was related to their Alps metal deposits running out.

AWood said...

@zardos

I agree with you on a few things. Most definitely the northerners were more primitive, at least in the sense their equipment was more rudimentary and bronze needed to be imported, where as the southerners were probably manufacturing it themselves. I think the southern army might have been an early offshoot of Halstatt A, and the only reason for the southerners to be here would be to get amber to decorate their stuff. Maybe this was the root cause of the battle, if trade went awry, or some political backstabbing, who knows. I would anticipate the wide range of men who seem to span west-central Europe kind of mirrors Bell Beaker networks, and I predict they formed one side of the battle. None of the "outliers" are I2, so I suspect they were from the local side, but this is just a guess. If I'm not mistaken the R1a guy plots with Baltic CWC, so maybe he was part of a third group or the locals?

Someone may have alluded to the fact the winning side may have buried or burned their dead leaving very little trace, and all these dead men belonged to the losing side. The lone R1a outlier might give weight to that possibility, and the local I2 guys were on the side of the southerners? We need to see more than 17 remains to shed more light on this, but it's all very interesting.

Anthony Hanken said...

Off topic but there were a couple new abstracts released.

Genetic continuity between ancient and recent populations in Central Asia

Perle Guarino-Vignon 1, Céline Bon 1, Nina Marchi 1 2, Evelyne Heyer1

1 - UMR 7206, Muséum National d’Histoire Naturelle, Paris, France · 2 - CMPG, Institute of Ecology and Evolution, Universityof Berne, Berne, Switzerland.

Southern Central Asia has been a crossroad for the movements of populations, cultures and goods between Europe, South Asia,East Asia and Middle East since prehistory. It has been under the influence from the North with the Steppe cultures and from the South with the Iranian culture for millenia. During the last 2000 years, populations speaking Turco-Mongol languages migrating from the eastern Asian populations partially replaced Indo-Iranian speakers. In precedent studies focusing on modern populations,it has been showed that there is a strong genetic difference between the modern Indo-Iranian population, and the Turco-Mongol speakers, with various amounts of admixture [1]. However, little is known about the origin of Indo-Iranian populations before 2000 years BP. Based on archaeological data, it has been suggested that Iron Age populations in Southern Central Asia were both related to Central Asia steppe populations (such as Andronovo) and Iranian population [2]. Despite its rich and complex history,Central Asia lacks genetic studies involving genomes from well-characterized and defined modern populations and ancient DNA to unravel the peopling of this region.

Samuel Andrews said...

@zardos,

The thing is there is no example of the LP allele in Neolithic farmers. For all we know Corded Ware got the allele from their Steppe ancestry.

If Corded Ware got the LP allele from a farmer population, that wouldn't change that Corded Ware IEs are the ones who made the allele widespread.

Anthony Hanken said...


Metagenomes and ancient human lineages from a pre-LGM layer of Satsurblia cave in the Caucasus

Pere Gelabert1,10, Susanna Sawyer1,10Anders Bergström2,10, Thomas C. Collin3, Tengiz Meshveliani4, Anna Belfer-Cohen5, David Lordkipanidze4, Nino Jakeli4, Zinovi Matskevich6, Guy Bar-Oz7, Daniel M. Fernandes1, Olivia Cheronet1,Kadir T. Özdoğan1, Victoria Oberreiter1, Mareike Stahlschmidt9, Pontus Skoglund2,11, Ron Pinhasi1,11

1 - Department of Anthropology, University of Vienna, Vienna, Austria · 2 - Ancient Genomics Laboratory, Francis Crick Institute, London, UK · 3 - School of Medicine & Medical Science, University College Dublin, Dublin, Ireland · 4 - Georgian State Museum, Department of Prehistory, Tbilisi, Georgia · 5 - Institute of Archaeology, Hebrew University, Jerusalem, Israel · 6 -Israel Antiquities Authority, Jerusalem, Israel · 7 - Zinman Institute of Archaeology, University of Haifa, Haifa, Israel · 9 -Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany · 10 - These Authors Contributed Equally · 11 - These Authors Contributed Equally.

Recent studies recovered multiple mitogenomes of various Homo lineages and other mammals from Late and Middle Pleistocene from archaeological sediments [1]. This approach opens a new horizon for the study of human evolution, admixture history and dispersals as either a complementary or an alternative approach to paleogenomic studies of human remains. It also holds the potential to provide new information about past environments, and human subsistence and behaviour.

Here we present genomic data from sediment from an Upper Paleolithic occupational level of Satsurblia cave, Imereti region,western Georgia, dated to 25, 000 years before present (bp). The site has yielded a rich archaeological record for Upper Palaeolithic occupation which spans from 33,000-14,000 years before present [2]. A previous study of a complete human genome from a human right temporal bone from this site, dated to the Late Upper Palaeolithic (13,132–13,380 cal bp) [3], showed that the post-LGM population which inhabited this region, were ‘Caucasus hunter-gatherers’ (CHG), a main source population for several Eurasian populations. However, the genetic composition of the pre-LGM inhabitants of the region remains unknown as there are no genomic data for this period for Western Eurasia.

Matt said...

Using the data I have for samples in G25, which is only a few of them (may not be the most current or correct I'd caution), then putting onto the Vahaduo West Eurasia/Europe PCA and adding onto them the y-dna assignments, from broadest to narrowest focus: https://imgur.com/a/oWycaD2

Not much more that I can say that hasn't been said though. In summary, in West Eurasian PCA, the samples seem to fit on a cline that parallels the present day European cline, but is shifted towards WHG / "West". The order of the samples goes such that the R1a->I2->R1b in "North"->"South". Two R1b samples can fit on the cline from CWC_Baltic_Early/Yamnaya->MN Europe. Then in North European PCA, none of the samples are really too strongly Western European in character whether R1b or not; sample WEZ35 overlaps Czechs and Hungarians, then even the sample WEZ57 which is Basque like in West Eurasia PCA is a bit shifted in position towards present day Eastern Europe relative to what a present day Basque sample is like in North Europe PCA.

Samuel Andrews said...

WEZ57=R1b, France
WEZ54=R1b, Spain.
WEz56=R1a, East Europe.

From main Welzin cluster there are five samples in G25 PCA.

Three cluster close together could be same population:
WEZ58: I2a2a
WEZ15: I2a2a
WEZ59: R1b

Two samples are basically the same but cluster signifcantly west of them. Maybe a different population from same region.
WEZ61: No Y DNA
WEZ35: R1b

It seems maybe R1b samples are roughly from same region as I2a samples. Maybe from same population.

Archi said...

@Romulus

"The losers were Urnfield, who we know carry I2a2 and come from an area rich in P312."

Do not talk nonsense. In the first half of the 13th century BC. Urnfield culture didn't even exist.

Don't fantasize, we only know the DNA of the Urnfield culture from one cave and there are completely different haplogroups.

Bronze Urnfield Germany Lichtenstein Cave, near Dorste, Lower Saxony [M1, M2, M7] 1000 BC M I2a2b
Bronze Urnfield Germany Lichtenstein Cave [M3, M6] 1000 BC M I2a2b
Bronze Urnfield Germany Lichtenstein Cave [M14] 1000 BC M I2a2b?
Bronze Urnfield Germany Lichtenstein Cave [M4, M5, M19] 1000 BC M I2a2b
Bronze Urnfield Germany Lichtenstein Cave [M8, M16] 1000 BC M I2a2b
Bronze Urnfield Germany Lichtenstein Cave [M10] 1000 BC M R1a1?
Bronze Urnfield Germany Lichtenstein Cave [M11] 1000 BC M R1a1?
Bronze Urnfield Germany Lichtenstein Cave [M9] 1000 BC M R1b

Bronze Urnfield / Lusitanian Germany Halberstadt [I0099 / HAL 36] 1193-979 calBCE (2889±30 BP, MAMS-21484) M R1a1a1b1a2 (Z280)

@AWood

"I think the southern army might have been an early offshoot of Halstatt A"

Too early for Hallstatt A. As far as I understand it is definitely not Hallstatt A.

----------------

n general, everything is bad with this Tollense battle. For 25 years of excavations, not a single book has been published, except for a few tiny articles on particular issues, nothing has been published, not even anthropological research has been done, although it is much faster to do it than to do genetic research. At this rate, we will understand something there in 250 years.

Rob said...

The difference in weaponry just reflects Internal ranks . There’s was a mounted officer class then the rank & file with whatever they could muster (eg clubs, axes).

Rob said...

Vatya culture in hungry was mentioned
That current sample (from Allentoft ‘15) falls under the same I2a2 as these guys.

zardos said...

There was a lot of research, but so far only pieces were published, still many uncertainties. The big publications and conclusions are still not there.
But I really think that alliances fought and most of the dead are supposed to be from the losing side. In the articles linked they mention groups of well-equipped warrior burials from the region and same time. Some compared them to some kind of brotherhood. To get and compare them with the Tollense remains could be enlightening.
Since archaeologists wrote about that, this should be the way to go.

Samuel Andrews said...

A read a few things on Tollense battle.

It sounded like the bodies were not buried. And by luck fell into a river and were preserved.

Why are you guys referring to burials? I must have read wrong.

zardos said...

Some even died in the water, so they were not robbed by the winners.

I spoke of nearby warrior burials which might represent true locals. Not from the river, just nearby.
And thats the only way to be sure which warriors at Tollense were locals and which not, by testing local burials too.

Samuel Andrews said...

@zardos,

Well, the ancestry of majority of published Tollense soldiers matches Bronze age Hungary. They look like they are from the Carpthian Basin. But of course we don't understand Bronze age regional variation well yet. None the less, they don't look like descendants of Corded Ware, Bell beaker, or Unetice even though many carry R1b. So, I tend to think they are not local to Northcentral Europe.

Halberstedt_LBA is from Northeast Germany, dates 1000 BC, is not like Tollense Soldiers.

Samuel Andrews said...

@zardos,

It would be cool to see DNA from nearby warrior burials. This would be the next step in understanding the Tollense battle.

It is surprising flint arrows were still being used as late as 1200 BC. Do local burials in Northern Europe from 1200 BC include flint weapons?

Davidski said...

@All

If you want these new Tollense samples in the Global25, then request the genotype data from these guys.

jburger@uni-mainz.de

daniel.wegmann@unifr.ch

zardos said...

@Sam: "Well, the ancestry of majority of published Tollense soldiers matches Bronze age Hungary. They look like they are from the Carpthian Basin. But of course we don't understand Bronze age regional variation well yet. None the less, they don't look like descendants of Corded Ware, Bell beaker, or Unetice even though many carry R1b. So, I tend to think they are not local to Northcentral Europe."

The general analysis points to a region like Southern Germany and Bohemia as a good fit, with relations up to the North in Poland and Germany. Actually pretty much to the border zone with the next, more Northern sphere. They were related to the Pannonian-Carpathian sphere, but not directly from there, at least there is little which would prove it or make it necessary.

Its just they were not Proto-Germanic, but a different people, probably closest to Celts paternally, but without being some, but from a now extinct IE or even non-IE group. What their opponents were, which took away most of their dead, is even more of a mystery.

Did the single R1a guy stick out autosomally by the way?

"It is surprising flint arrows were still being used as late as 1200 BC"

No its not, because arrowheads get lost easily and are fairly expensive, while the stone ones are still effective. So for poor people, they were just first choice. And that they were effective you can see, beause varous of the diseased have flint arrowheads in or at their bones, suggesting they were wounded or killed by them.
And then again, the elaborated bronze arrowheads found (Tüllenpfeilspitzen) with a cup are very rare up in the North, unknown in Scandinavia. The same is true for many more goods found, all point to Southern Central Europe.

You can read on the issue this article with the fitting title:

https://www.academia.edu/11604957/Not_at_all_obsolete_The_use_of_flint_in_the_Bronze_Age_Netherlands

ambron said...

R1a has a large Balto-Slavic drift. Lucas groups him with northeastern Poland.

zardos said...

Now I see that WEZ56 was the outlier and now being confirmed R1a, being pulled heavily to the North East. Well, he might be a warrior which participated on the campaign more or less by chance or he is a good representative of a stronger element among the winners, which carried most of their dead on dry ground away...

Samuel Andrews said...

I emailed the authors. We'll see if they make the genotype data public.

It is frustrating when researchers don't understand the power of ancient DNA has for understanding history.

"All the soldiers are generally genetically similar, therefore they all come from the same exact population. Nothing more to look for."

Parastais said...

Is that WEZ56 similar to (somewhat) contemporary Baltic BA samples or more similar to modern Slavs or Balts?

ambron said...

Two samples from Kuyavia (N47 and N49, both I2a2), one more autosomal similar to WEZ56 and the other to the main Welzin cluster, show that in the Pomeranian region in the old days people with widely different autosomal diversity could live side by side. WEZ56 does not have to be an outliner, because we also have R1a-Z283, Z280 from Halberstadt from the nearby area and period.

Tigran said...

So what caused the selection for LP? A switch to milk from cheese. Were milk and yogurt consumed in the European neolithic or was most dairy cheese?

Samuel Andrews said...

Exactly. That is the job of archaeologists. Obviously, 2000-500 BC, there was dependence on dairy in diet in much of EUrope.

In South Asia, the only populations in which the LP allele is popular is in ones traditionally dependent in dairy which is interesting and no coincidence.

Archi said...

For that time, we can only talk about three large opposing formations, the Tumulus culture, the Proto-Lusatian culture and the Nordic Bronze Age culture. All of them were hostile to each other, so on which side the Lusatians were playing was not at all determined.

I must say that we really do not know at all the autosomal distribution between the cultures of that era, therefore, it is now impossible to assert who is from which region.

zardos said...

@Tigran: Milk was consumed much earlier, without LP spreading fast. Its no all or nothing thing, like drinking milk or not. The shift must have been caused by an extreme consumption of and dependence from milk. And it helped the non-LP people too, because the rates were so low, it spread among non-tolerant populations. It was just that among these heavy milk users, those with LP got the edge, less problems, more energy, less likely to get sick from it - and I don't mean the direct, but probably also indirect effects of intolerance, like being more vulnerable to infections of the digestive tract or something like that.

So the real question is, when and why did people start, which people and where, to become so completely dependent from fresh milk. We know a similar case from the Masaai, which even live primarily from the milk, meat and blood of their cattle.
https://www.wired.com/2012/09/milk-meat-and-blood-how-diet-drives-natural-selection-in-the-maasai/

Their diet is extremely rich in proteins and fat. Interestingly, there is another allel spreading almost parellel, which reduces blood cholesterol and of course there is depigmentation, happening parallel too.
I think that all these traits, and possible others, so far undetected, being connected by a change of the selective regime, related to the diet and way of life of the people affected.

So it was more than just "drinking milk occasionally", there is no way to get that kind of selective pressure like that. It was a complete orientation on animal products. Logically, this fits better into nomadic or half-nomadic steppe people, but that's not the case it seems. There are heavy milk consumers on the steppe, to this day, which are largely lactose intolerant.

It must have been quite an extreme change and dependency, probably caused by worsening living conditions at one point, leading to adaptations which became very successful afterwards and spread with the people profiting from it.

Tigran said...

@zardos

Thanks.

Do you know if yogurt existed back then or is that a recent Central Asian Turkish thing?

Samuel Andrews said...

@zardos,

Interesting. Are Europeans traditionally more a diary culture than other parts of the world? I would think this could be a cultural-significant thing for Europeans going back to Bronze age.

Milk is as standard as water in America but only because of industrial farming in last 100 years.

I read that before then milk was luxury. But, I don't believe that.

zardos said...

@Samuel: It became a luxury for some workers and slaves in America, but it was a common food throughout most of Europe from earliest times recorded. Many families survived times of famine and catastrophy only relying on the milk of a cow or goat. Especially Germanics seem to have relied heavily on milk and meat. If being poor cereals and milk was still a good meal. So especially the poor people are supposed to have dependent from milk and milk products primarily. But the poor were the majority, so...

@Tigran: I think some predecessor of Yoghurt was used much earlier, like some forms of cheese, but that's so remarkable about it. Milk products were used intensively before, so it must have been a new and drastic increase. Like the poor Germanics I referred too, for which milk became THE nutrient food they had to rely upon, many times daily probably, in high amounts. Being poor, bad nutrition, relying upon milk, then getting sick = end. So every advantage counted.

Onur Dincer said...

@zardos

So it was more than just "drinking milk occasionally", there is no way to get that kind of selective pressure like that. It was a complete orientation on animal products. Logically, this fits better into nomadic or half-nomadic steppe people, but that's not the case it seems. There are heavy milk consumers on the steppe, to this day, which are largely lactose intolerant.

If you mean the Turco-Mongol nomadic peoples of the steppe, they are not heavy milk consumers but heavy consumers of fermented milk products like kumis, yogurt and cheese, which are all low in lactose due to the fermentation process in their production. It is still possible that the earlier IE steppe nomads of the steppe were heavy milk consumers.

matthayichen said...

My theory of LP spread goes like this. There is a cultural inflection point where selection really takes off. It is the point where the cultural wisdom goes from "don't drink raw milk or you'll get sick" to "you have to drink fresh milk to be big and strong".

Perhaps migration and founder effects can trigger this cultural change locally. And wherever such (conquering) clans are dominant, everyone else will try to follow the cultural norms or emulate the elites and those who get sick will simply be considered inferior stock and naturally sickly, and weeded out.

Just my two cents as a potential alternative to the 'heavy dependency on raw milk, yoghurt not an option' theory.

Davidski said...

@matthayichen

And wherever such (conquering) clans are dominant, everyone else will try to follow the cultural norms or emulate the elites and those who get sick will simply be considered inferior stock and naturally sickly, and weeded out.

Yes, interesting theory. But I suspect it's too hardcore for the Germans to have even considered it an option in a formal paper.

Samuel Andrews said...

When I read, matthayichen's quote it looked like the way a Marxist archaeologist would describe the likely origin of the LP allele. Yet Davidski, thinks it looks Nazi(ish)/Fascist hence German researchers would never touch such an idea.

Interesting, how both ideologies have a similar dark view of human nature.

Kind of like, how archaeologist see a rich ancient burial as an indication of something negative: "Social Inequality." Instead, of the positive: a very cool guy