"The Genetic History of the Southern Arc: A Bridge between West Asia & Europe"

[Bruzmi]

The samples we've got so far from Montenegro, Serbia and Albania tell a different story. I personally didn't expect we'd see so much structure in the Medieval Balkans but here we are. Those Montenegrin samples that are right on the border with Albania and are way more Slavic than modern Montenegrins are a smoking gun. And before you get defensive again, we don't know the details of how it happened. It could have involved intermediate, bilingual groups, it could have involved nomadic people like Vlachs and it could be a long-term process of small-scale admixture that lasted centuries. My pet theory is that the big homogenisation that resulted in the modern cline was facilitated by the Rum millet reforms of the Ottoman Empire. Anyway, I hope in the future we get a big Byzantine and Ottoman aDNA study that will illuminate these processes.

Yes, the 11th century sample from Montenegro is ~15%+ more Slavic-like than modern Montenegrins. He is J-L283 so in terms of Y-DNA there's not much more to say. My opinion about Montenegro is that the modern profile is largely the result of Albanian clans who via common religion gradually became part of the Slavic population in the late Middle Ages and via massive founder effects fundamentally altered Y-DNA distribution in Montenegro today. Montenegro is so well tested that we can observe these processes in a clear timeline.

In terms of Y-DNA, there's very little early contact between Albanians and the "north-western" South Slavs (ancestors of Croats, Serbs, Bosniaks, Montenegrins). I-PH908, the characteristic clade of these groups, represents 23/1463 (~1.60%) of samples on Rrënjët. It may be up to 2% in total since quite a few who should be I-PH908+ haven't been tested beyond I-Y3120. Early contact situations favor more "south-eastern" groups which carried I-Z17855 or I-Y18331 and it's under these clades that the majority of Albanian I-Y3120 would be placed if they tested further.

[DFSTFD]

The profile of modern like Northern Italian genetics was already in place during the bronze age. They were less homogenous, but the averages were already there. This is corroborated by northern adriatic samples from IA croatia and slovenia which looked identical to a N. Italian profile and emerged from the commonly accepted balkan route of italic langauge migrations into italy. The Etruscans/latins forming a cline with such genetics is no coincidence either. What's more is we already see the same profiles in Roman military camps abroad during the early empire, and you see them in C. Italy more commonly to the tune of about 20% of the population by the late empire. Those who believe northern italian like genetics were birthed from some germanic or frankish mixing event during the middle ages/late empire are setting themselves up for massive disappointment. Modern Northern Italians have far too much neolithic anatolian ancestry to be a result of central or northern european influence.

This is a bit off-topic but this doesn't seem to be exactly the case.

Comparing the varied north Italian set in G25 (an important caveat that some might not be fully representative of a certain area, some might have overrepresentation from more marginal regions like the Aosta Valley set and the Northeast set that seem to have their own recent/separate admixtures, more clear in the Northeast's Slavic-related input, so it might influence the impression in both directions; basically we could do with an even bigger set for some north Italian regions) with both the directly geographically adjacent IA populations (France, Iberia, Slovenia, central Italy) and the whole Italian BA-Neolithic set we currently have, north Italy seems shifted towards the 'south'. Lumped together and separately with all Italian groups.

I added the Campiglian CEU Etruscan outlier as well because it's a radiocarbon dated 7th century BCE sample so relatively early even though his ultimate geographic associations are uncertain.

This implies at least eastern imperial input of some kind. But I agree with the last part of your comment.

The ideal of course is to see what various parts of north Italy actually look like in the IA, rather than guess contextually, and compare to that.

Yes, but what the paper says is that the basic components have comparable proportions from BA to now. One model explains this by saying there is little to no admixture after BA, this obviously works. Another model says that in antiquity/Early middle ages there was admixture that would have increased CHG/ANF and then a second admixture that would have increased EHG, resulting in comparable proportions as before Antiquity. This also works. So I don't see how the statement from the paper invalidates either view.

Also perhaps a bit off-topic, though at least the broad Balkan changes throughout some periods have been brought up in the paper, but my two cents.

Separate from the specifics, which have been discussed a lot and obviously can change quite a bit depending on the specific model so there will be some disagreement between people, in the broad sense we can tell the difference in G25 (and arguably with more proximal modelling in qpAdm, per the Olalde et al. paper, unlike the paper's distal one in this case). The old Balkan cline follows a somewhat different path. The modern one is more "eastern-shifted" both in West Eurasia PCA terms (towards the ANE-rich direction, i.e. higher combined EHG+CHG, lower Anatolia_N) and Europe PCA terms (more towards the combined Baltic+Aegean direction, less towards the Sardinian). We can certainly tell something changed in finer scale (and arguably more broadly as well but you see the difference better in within Europe PCA in some cases) but the proportions are much more arguable depending on your choices.

Another outlier from Montenegro Iron Age, shifts North-East

Distance to: MNE_IA:I13170

This particular individual isn't radiocarbon dated and doesn't look like what we see with confirmed BA-IA samples. It's probably a similar case of misdating as some of the misdated Bezdanjaca individuals that might be even modern. Funnily enough this one plots very close to the Asia Minor Slavic individual TUR_Marmara_Ilipinar_Byz3.

[Karagjoz]

Yes, the 11th century sample from Montenegro is ~15%+ more Slavic-like than modern Montenegrins. He is J-L283 so in terms of Y-DNA there's not much more to say. My opinion about Montenegro is that the modern profile is largely the result of Albanian clans who via common religion gradually became part of the Slavic population in the late Middle Ages and via massive founder effects fundamentally altered Y-DNA distribution in Montenegro today. Montenegro is so well tested that we can observe these processes in a clear timeline.

In terms of Y-DNA, there's very little early contact between Albanians and the "north-western" South Slavs (ancestors of Croats, Serbs, Bosniaks, Montenegrins). I-PH908, the characteristic clade of these groups, represents 23/1463 (~1.60%) of samples on Rrënjët. It may be up to 2% in total since quite a few who should be I-PH908+ haven't been tested beyond I-Y3120. Early contact situations favor more "south-eastern" groups which carried I-Z17855 or I-Y18331 and it's under these clades that the majority of Albanian I-Y3120 would be placed if they tested further.

Yes, it looks like the early medieval slavic input into albanians originated mostly from the region of ukraine and romania though a small fraction may have come from the north at the border zones (mostly as ph908)

Interesting that the montenegrin with very slavic autosomal carried j2b l283, points to a big slavic presence in montenegro since at least 10th century and likely an earlier assimilation of this j2b line

[Kale]

It would be nice to see if the Baltic Corded Ware samples that showed no farmer admixture whatsoever have it too. Plinkaigalis242 and Gyvakarai1 from Mittnik 2018 and Latvia_LN1 from Jones 2017. Corded Ware may behave differently from Yamnaya.

https://eurogenes.blogspot.com/2018/...m-genetic.html

Also would be nice to look how X to autosomal ratio for CHG is in these samples.

Qpadm has a hard time telling apart ANF and Levant_N in such small quantities, they are just too similar. But if you use Progress_Eneolithic as a baseline, steppe populations need a sizeable chunk of WHG.
Great 2-way model here.
Afanasievo_KarasukIII.SG
RUS_Stavropol_Progress_Eneolithic 0.886518 0.0216123 41.0192
Romania_GuraBaciului_CA.SG 0.113482 0.0216123 5.25080
Tail: 0.50
right = c('Congo_Mbuti.DG', 'Anatolia_Boncuklu_N.SG', 'Iran_Wezmeh_N.SG', 'CHG.SG', 'Levant_N', 'Taforalt', 'Bichon.SG', 'Italy_GrottaContinenza_HG.SG', 'RUS_Arkhangelsk_HG.SG', 'Botai.SG', 'Andaman_100BP.SG', 'RUS_Primorsky_DevilsCave_N.SG', 'Peru_RioUncallane_1800BP.SG')

[Gentica277282]

Qpadm has a hard time telling apart ANF and Levant_N in such small quantities, they are just too similar. But if you use Progress_Eneolithic as a baseline, steppe populations need a sizeable chunk of WHG.
Great 2-way model here.
Afanasievo_KarasukIII.SG
RUS_Stavropol_Progress_Eneolithic 0.886518 0.0216123 41.0192
Romania_GuraBaciului_CA.SG 0.113482 0.0216123 5.25080
Tail: 0.50
right = c('Congo_Mbuti.DG', 'Anatolia_Boncuklu_N.SG', 'Iran_Wezmeh_N.SG', 'CHG.SG', 'Levant_N', 'Taforalt', 'Bichon.SG', 'Italy_GrottaContinenza_HG.SG', 'RUS_Arkhangelsk_HG.SG', 'Botai.SG', 'Andaman_100BP.SG', 'RUS_Primorsky_DevilsCave_N.SG', 'Peru_RioUncallane_1800BP.SG')

I would assume because ppnb has heavy Anatolian admixture

[DFSTFD]

Qpadm has a hard time telling apart ANF and Levant_N in such small quantities, they are just too similar. But if you use Progress_Eneolithic as a baseline, steppe populations need a sizeable chunk of WHG.
Great 2-way model here.
Afanasievo_KarasukIII.SG
RUS_Stavropol_Progress_Eneolithic 0.886518 0.0216123 41.0192
Romania_GuraBaciului_CA.SG 0.113482 0.0216123 5.25080
Tail: 0.50
right = c('Congo_Mbuti.DG', 'Anatolia_Boncuklu_N.SG', 'Iran_Wezmeh_N.SG', 'CHG.SG', 'Levant_N', 'Taforalt', 'Bichon.SG', 'Italy_GrottaContinenza_HG.SG', 'RUS_Arkhangelsk_HG.SG', 'Botai.SG', 'Andaman_100BP.SG', 'RUS_Primorsky_DevilsCave_N.SG', 'Peru_RioUncallane_1800BP.SG')

GuraBaciului is the extremely HG-rich GB1 individual. Makes sense if you consider the Progress end of the Don-Volga steppe. Have you tried/gotten any good models with just Khvalynsk, both the average and the individuals (since they vary quite a bit horizontally on the PCA) or with the combination of Khvalynsk+Progress?

[Kale]

I would assume because ppnb has heavy Anatolian admixture

I suspect there may have been a cline of AHG > ANA in the Eastern Mediterranean. Then parallel process of Mesopotamian farmers effecting both Anatolia and the Levant. Then like you say actual Neolithic Anatolian ancestry in the Levant.

[Ahuwarhd]

This is a bit off-topic but this doesn't seem to be exactly the case.

Comparing the varied north Italian set in G25 (an important caveat that some might not be fully representative of a certain area, some might have overrepresentation from more marginal regions like the Aosta Valley set and the Northeast set that seem to have their own recent/separate admixtures, more clear in the Northeast's Slavic-related input, so it might influence the impression in both directions; basically we could do with an even bigger set for some north Italian regions) with both the directly geographically adjacent IA populations (France, Iberia, Slovenia, central Italy) and the whole Italian BA-Neolithic set we currently have, north Italy seems shifted towards the 'south'. Lumped together and separately with all Italian groups.

North east and aosta valley Italians do not appear slavic or celtic related at all as they seem to cluster relatively tightly with the rest of po valley. To me they look to be a continuation of the bronze age established continuum in northern italy and the fact that 7th century PCA results from Aosta valley overlap moderns corroborates this idea. I think you're simply reading too much into slight regional differences that have likely existed since the pre-roman era.

As far as modelling BA northern italian averages on a PCA via G25 - this is impossible to do atm because we're missing 2 of the 5 sample coordinates that exist from broion BA. Of the remaining 3 that do exist as far as G25 coordinates go, they seem to be least steppe impacted samples, meaning the average on G25 is currently drastically skewed towards sardinia, in contrast to what the study reported. If you were to add in the 2 remaining high-steppe samples the average would look extremely modern to N. Italy - even more so than IA etruscan/latin samples.

This implies at least eastern imperial input of some kind. But I agree with the last part of your comment.

I don't think Roman era E med like genetics will have much of an impact on N. Italians at all to be honest. If they do it would have to be exceedingly minor. To me, N. Italians today seem to be a holdover from a very clear genetic distribution that emerged out of the northern Balkans during the bronze age and early iron age. These people were not impoverished in CHG related ancestry and don't require additional sums of it from east meds. Even the lowest steppe impacted samples of broion had already ~%10 CHG related ancestry. We can most definitely assume the higher impacted steppe samples would have significantly more.

But I agree with the last part of your comment.

Glad we see eye to eye there. I just don't see how it's even possible for anyone to get N. italian average out of any type of cross admixture between E. Med/S.Italian and C. Europe. It would require Northern Italians to have nearly 10% less barcin than they actually have today and also a good bit more WHG which they nearly totally lack. You couldn't even get N. Italian averages out of a cross between etruscan IA and CEU, because the CHG related ancestry would be too low and the WHG still too high. The N. Italian autosomal profile is very old, and if it weren't for its longstanding genetic introgressions into the rest of post roman Italy, it would effectively be an island of its own, today.

The ideal of course is to see what various parts of north Italy actually look like in the IA, rather than guess contextually, and compare to that.

I totally agree. It's a shame that there are no iron age studies going on in Po valley or its overlooking foothill valleys.

[DFSTFD]

North east and aosta valley Italians do not appear slavic or celtic related at all as they seem to cluster relatively tightly with the rest of po valley. To me they look to be a continuation of the bronze age established continuum in northern italy and the fact that 7th century PCA results from Aosta valley overlap moderns corroborates this idea. I think you're simply reading too much into slight regional differences that have likely existed since the pre-roman era.

Half of the individuals in the particular "Northeast" set compared to the other half are visibly off-cline compared to the rest of north Italy towards Balkan-like groups and score Baltic-rich at higher levels. They must have "recent" Slavic-related admixture, though I assume they come from border areas and are less representative. Those outliers to the general Italian cline is what I'm referring to, not the whole set.

I don't think Roman era E med like genetics will have much of an impact on N. Italians at all to be honest. If they do it would have to be exceedingly minor. To me, N. Italians today seem to be a holdover from a very clear genetic distribution that emerged out of the northern Balkans during the bronze age and early iron age. These people were not impoverished in CHG related ancestry and don't require additional sums of it from east meds. Even the lowest steppe impacted samples of broion had already ~%10 CHG related ancestry. We can most definitely assume the higher impacted steppe samples would have significantly more.

Not "east med" like necessarily but they might be Balkan-Pannonian related as alexfritz has suggested before, for example; but something from the "eastern empire" nonetheless. The north Italian centroid is close to HRV, though that might be to an extent due to overrepresentation of more southern individuals within it, i.e. off cline relative to any of the previous Beaker/Italic-related groups we at least currently have in G25 or compared to any direct neighbors i.e. whether you draw a cline from IA Slovenia to France to Iberia to central Italy. Though the Adriatic coast of central Italy has furnished us that HRV-like individual of likely recenty Balkan origins so the interplay in the eastern half of at least central Italy will be interesting to see.

[Kale]

GuraBaciului is the extremely HG-rich GB1 individual. Makes sense if you consider the Progress end of the Don-Volga steppe. Have you tried/gotten any good models with just Khvalynsk, both the average and the individuals (since they vary quite a bit horizontally on the PCA) or with the combination of Khvalynsk+Progress?

- Progress + Romania + Khvalynsk = doesn't want the latter
- Progress + Barcin + Khvalynsk = fail (not enough WHG)
- Progress + Romania + Khvalynsk + Maikop = doesn't want either of the latter
- Khvalynsk + Barcin + IronGates_HG + Maikop = -20% Barcin
- Khvalynsk + Barcin + IronGates + Caucasus_Eneolithic = fail, -5% ANF, 4% IGHG.
- Progress + Maikop + Ukraine_Dereivka_NHG = tail 0.10, proportions 81/6/13
- Progress + Ukraine_Dereivka_NHG = tail 0.06 proportions 88/12

Progress + Steppe-adjacent (thus reasonably HG-rich) farmers seems the best bet.