Population History of Sardinia

Angela

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Chiang et al:
"Population history of the Sardinian people inferred from whole-genome sequencing"

See:
http://biorxiv.org/content/early/2016/12/07/092148

"The population of the Mediterranean island of Sardinia has made important contributions to genome-wide association studies of traits and diseases. The history of the Sardinian population has also been the focus of much research, and in recent ancient DNA (aDNA) studies, Sardinia has provided unique insight into the peopling of Europe and the spread of agriculture. In this study, we analyze whole-genome sequences of 3,514 Sardinians to address hypotheses regarding the founding of Sardinia and its relation to the peopling of Europe, including examining fine-scale substructure, population size history, and signals of admixture. We find the population of the mountainous Gennargentu region shows elevated genetic isolation with higher levels of ancestry associated with mainland Neolithic farmers and depleted ancestry associated with more recent Bronze Age Steppe migrations on the mainland. Notably, the Gennargentu region also has elevated levels of pre-Neolithic hunter-gatherer ancestry and increased affinity to Basque populations. Further, allele sharing with pre-Neolithic and Neolithic mainland populations is larger on the X chromosome compared to the autosome, providing evidence for a sex-biased demographic history in Sardinia. These results give new insight to the demography of ancestral Sardinians and help further the understanding of sharing of disease risk alleles between Sardinia and mainland populations."

Nice to get confirmation from whole genomes for what some of us have been saying for years, i.e. that there is structure in Sardinia.

I'm less sure what they mean about this hunter-gatherer ancestry in the isolated mountain areas. The "consensus" has been that the signs of European pre-Neolithic hunter-gatherers in the archaeology was from occasional visits, not long term settlement which would have significantly impacted the genetics of the island (although I argued against that at one time).

Lately, I've been of the opinion that the "Neolithic" substratum is actually from migration from Old Europe.

By the way, they're basically talking about the area around Ogliastra.

https://www.youtube.com/watch?v=NOegu2jgJcY



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Some interesting bits:

" PCA shows a one-dimensional isolation-by-distance configuration around the Mediterranean, from North Africa through the Near East and then towards Iberia (Novembre and Stephens 2008, Henn et al. 2012, Botigue et al. 2013, Paschou et al. 2014) (Figure 4A,). The effective migration surface shows the Mediterranean Sea as having low effective migration (Figure 4B), isolating Sardinia"from neighboring mainland populations, with stronger isolation between Sardinia and North Africa than mainland Europe (Figure 4B).

I tried to explain how difficult it is to go directly from the mainland to Sardinia and even more so from North Africa to Sardinia because of the air and sea currents, but it was heavy going. Maybe women are presumed to know nothing about navigation. Now here is the proof in genetics.

"As an alternative visualization of pan-Mediterranean population structure, an analysis using the ADMIXTURE software inferred four ancestral components, with one component associated primarily with Sardinians and Southern Europeans (“red”), and remaining components corresponding to North African (“blue”), Middle East and Caucasus (“purple”), and Northern Europeans (“green”) (Figure 4C; see Figure S3 for results at other values of K). The Arzana individuals contained 100% of this red component and Sardinians from Cagliari contained 93% of this red component."

"e sharing with admixed Latino populations may reflect variants shared between Sardinian ancestors and ancestral European sources to admixed Latino populations (e.g. Iberian populations)."

"Due to its smaller long-term effective population size (Figure 5A), Sardinia is expected to have undergone accelerated rates of genetic drift. To correct for this when measuring similarity to other mainland populations, we used “shared drift” outgroup-f3 statistics (Raghavan et al. 2014), which are robust to population-specific drift. Using this metric, we find the Basque are the most similar to Sardinia, even more so than neighboring mainland Italian populations such as Tuscany and Bergamo (Figure S6A, S6B). This relationship is corroborated by identityby-descent (“IBD”) tract length sharing, where among mainland European populations, French Basque showed the highest median length of shared segments (1.525 cM) with Arzana."

We also tested the affinity between Sardinians and Basque with the D-statistics of the form D(Outgroup, Sardinia; Bergamo or Tuscan, Basque)...We find that Sardinia consistently showed increased sharing with the Basque populations compared to mainland Italians...In contrast, sharing with other Spanish samples in our dataset was generally weaker and not significant ( |Z| < 3.5; Figure S6C), suggesting the shared drift with the Basque is not mediated through Spanish ancestry."

This might be an indication that there were once French Basque like populations throughout southern France which then moved into Sardinia. While it's true, as I said, that it's difficult to reach Sardinia from the mainland, the route from southern France/northeast Spain to Sardinia isn't too bad. The other "relatively" easy trip is by sea from Greece, southern Italy, Sicily etc.

They do a much better job than Hellenthal at explaining any "recent" admixture in Sardinia. There is no indication of it in mainland, central Sardinia. One test failed to disclose any even in areas like Cagliari, the capitol, but one did find some.

"h previous reports (Moorjani et al. 2011, Loh et al. 2013, Hellenthal et al. 2014). For example, Cagliari individuals demonstrated ~7% of a non-Sardinian (“purple”) component that is found in substantial fraction among extant individuals from Southern Europe, Middle East, Caucasus, and North Africa. To assess this in detail we used the f3-test for admixture (Patterson et al. 2012) and found none of the Sardinian populations showed any evidence of admixture. In contrast, mainland Europeans, particularly Southern Europeans, show evidence of admixture from Near East and sub-Saharan Africa (Figure S8). Because f3-based tests for admixture may lose power when applied to populations that have experienced extensive drift post-admixture (Patterson et al. 2012), we also used a complementary LD-based approach (ALDER, Loh et al. 2013) to test each Sardinian population for admixture. Using this approach, a number of Sardinian populations, particularly those outside of Ogliastra, are inferred to be admixed (Table 1, Table S4). The inferred source populations are typically a mainland Eurasian population and a sub-Saharan African population. The admixture proportions range from 0.9% to 4% of sub-Saharan ancestry by the f4 ratio test (Patterson et al. 2012) (Table 1, Table S4) with estimated admixture dates of approximately 59-109 generations."

"found the best proxy for African admixture is sub-Saharan African populations, rather than Mediterranean North African populations, and we inferred the date of admixture as approximately 1,800-3,000 years ago (assuming 30 years per generation).

I differ with them in the following to the extent that I don't think that there was large scale migration from the Near East with the Arab invasions, although there was some.

"lack of a strong signal of North African autosomal admixture may be due to inadequate coverage of modern North African diversity in our reference sample. Alternatively, it may be due to a poor representation of ancestral North Africans. Present-day North African ancestry reflects large-scale recent gene flow during the Arab expansion (~1,400 years ago (Henn et al. 2012)). The sub-Saharan African admixture observed in the non-Ogliastra samples could be mediated through an early influx of migrants from North Africa prior to the Arab expansion, for example during the eras of trade relations and occupations from the Phoenicians, Carthaginians, and Romans (~700 B.C.- ~200 B.C.;"

"both ancient Neolithic farmer ancestry and pre-Neolithic ancestry are enriched in the Gennargentu-region. First, we find that shared drift with Neolithic farmers and with pre-Neolithic hunter-gatherers is significantly correlated with the proportion of “Gennargentu-region” ancestral component estimated from ADMIXTURE analysis, while shared drift with Steppe pastoralists has a weak negative correlation with Gennargentu-region ancestry (Figure 6B). Second, using supervised estimation of ancestry proportion based on aDNA (Haak et al. 2015), we estimate higher levels of Neolithic and preNeolithic ancestries in the Gennargentu region and higher levels of Steppe Pastoralist ancestry outside the region (Figure S10)"

"Intriguingly, on average, we find a higher proportion of the Gennargentu-region ancestry (“red” component in Figure S11) on the X-chromosome (37%) than on the autosome (30%, P < 1e-6 by permutation). As the Gennargentu-region ancestry is correlated with more ancient, Neolithic or pre-Neolithic ancestries rather than Bronze Age ancestries (Figure 6B), this finding suggests sex-biased processes in which more females than males carried the non-Steppe ancestries."

"These results demonstrate that sex-biased process occurred in the founding of Sardinia, which is above and beyond any putative sex-biased processes on the mainland."

I have some skepticism about this:
"Our analysis of crosscoalescent rates suggest the population lineage ancestral to modern-day Sardinia was effectively isolated from the mainland European populations approximately 330 generations ago. This estimate should be treated with caution, but corresponds to approximately 9,900 years ago assuming a generation time of 30 years and mutation rate of 1.25x10-8 per basepair per generation."

Here they address the point I raised about the uncertainty as to where the WHG ancestry was added:
. That pre-Neolithic hunter-gatherer and Neolithic farmer ancestry are enriched in this region of isolation suggests that the early populations of Sardinia were an admixture of the two ancestries, rather than the pre-Neolithic ancestry arriving via later migrations from the mainland. However, it remains to be seen whether this admixture principally occurred on the island or on the mainland prior to a Neolithic era influx to the island."

"The high frequency of particular Y-chromosome haplogroups (particularly I2a1a2 and R1b1a2) that are not commonly affiliated with Neolithic ancestry is one challenge to a model in which Sardinian principally has Neolithic ancestry."

I don't see the problem with the above. We already know of I2a1a2 samples from the mainland which are typically European farmer. They apparently represent absorbed male hunter-gatherer lines.
 
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There appear to be strong links between pre-Nuragic Sardinia and Southern France via the obsidian trade; it could all boil down to 'economics' in the end. Those trade networks and associations could, and probably did endure long enough to usher in a whole lot more--other interested parties, like "Beaker Folk."

Fanciful speculation, but I wonder if Arturo Issel was right about the Ligures.
 
Can we call them pureist Med people?
 
for the most part sardinians jus look like italians to me though, i cant see a difference really in Looks, but I dont know to many sardinian though to base that off of. I only think of Franco columbu the Body building compatriot of Arnold schwarzenneger when i think of sardinians :). and I wonder if previously sardianians had spoken a similar language to basque before they becae italicized, now that woud be interesing to prove.
 

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it's not that difficult to reach Sardinia from Central Italy through Corsica,; archaeology suggest that the Mesolithic, Cardial and part of the Bell Beakers settlers came there from Tuscany. In the supp info. Yamnaya ancestry runs from 0% in some Ogliastra villages to ~10% in the North and the South-West of the island.
 
No surprise the conservative results of the mountain area (Gennargentu, Nuoro), as it realy seems an island inside the island. But to take the Sardinian DNA as mostly unmixed i'm not so sure when looking at its history (Bell Beakers from the continent, shardana from the Aegean? lybi from Africa...).
 
for the most part sardinians jus look like italians to me though, i cant see a difference really in Looks, but I dont know to many sardinian though to base that off of. I only think of Franco columbu the Body building compatriot of Arnold schwarzenneger when i think of sardinians :). and I wonder if previously sardianians had spoken a similar language to basque before they becae italicized, now that woud be interesing to prove.

Italians have a lot of "Med" so obviously there's going to be some overlap, with some mainland Italians looking like Sardinians.

However, there's a lot of variety in Italy which just doesn't exist in Sardinia, in pigmentation for one thing, but also in body type, head type, facial features etc.

I just posted this video in another thread, but it also illustrates what I'm talking about. These are Italians too, and they don't look at all Sardinian.
https://www.youtube.com/watch?v=sBevfffdqVA

The man in the last picture I posted above is a type that would be unusual in Italy.

The people to whom the Sardinians are most related are the French Basques, and there is indeed a Basque look in some of them.

@Berun,
The point the authors are making is that those "other" influences did not penetrate into the internal Massif. Of course, all Sardinians are more related to one another than to anyone else.

@Bix,
I have great respect for Arturo Issel, but I think he was totally wrong about that, although given the Paleolithic finds in Liguria, I know what might have been affecting his opinion. No one in Europe is within the range of variation of even Mesolithic hunter-gatherers, much less the Paleolithic ones. EEF is a different story, which is the point of the paper. People overlapping with the EEF do still exist, in Sardinia.

@Cato,
Could you please provide some written sources for that as it pertains to Mesolithic and Cardial?
 
I have great respect for Arturo Issel, but I think he was totally wrong about that, although given the Paleolithic finds in Liguria, I know what might have been affecting his opinion. No one in Europe is within the range of variation of even Mesolithic hunter-gatherers, much less the Paleolithic ones. EEF is a different story, which is the point of the paper. People overlapping with the EEF do still exist, in Sardinia.

Certainly, I agree. I had something the authors said near the end of the discussion section in mind where they said:

"one might consider a model in which a yet unsampled sub-population during the Neolithic carried high proportions of the I2a1a haplogroup.

I think a population somewhere around the Rhone, or around the Gulf of Lion would be a good prospect. I read somewhere, but I can't recall where that ancient Ligures were far more widespread--they seem like an interesting lot, with some rather unorthodox sailing skills, vexing to the Romans. They were surely a well admixed with a long list of different groups by the time the Romans came around.

This subject has some genealogical pertinence to me, wondering how my particular brand of I-M26 got to the western Alps.
 
Certainly, I agree. I had something the authors said near the end of the discussion section in mind where they said:

"one might consider a model in which a yet unsampled sub-population during the Neolithic carried high proportions of the I2a1a haplogroup.

I think a population somewhere around the Rhone, or around the Gulf of Lion would be a good prospect. I read somewhere, but I can't recall where that ancient Ligures were far more widespread--they seem like an interesting lot, with some rather unorthodox sailing skills, vexing to the Romans. They were surely a well admixed with a long list of different groups by the time the Romans came around.

This subject has some genealogical pertinence to me, wondering how my particular brand of I-M26 got to the western Alps.

Sorry, Bix, I misunderstood. Yes, I too have read the theories where the Ligures were once a much more widely spread population, which goes along with the theory that they were an early group of Indo-European speakers. Certainly, they existed outside the boundaries of modern day Liguria, living, we are told, north of Massalia, and into northeastern Spain, although progressively more mixed with Iberians as they got further south. If they were Indo-European speakers they would have already been admixed by the time they were in Central Europe and would then have admixed with farmers again further south.

I too have a lot of interest in them, because depending on where you draw the lines, a lot of my ancestry comes from them. Unfortunately, all we have is some comments in ancient history, and some speculations based on a few bits of their language.

Have you taken a look at the Wiki article? It's not bad as a summary. I will say that the description of their character in the ancient sources rings true to me: sparely built and lean, but very strong physically and mentally. It was also said of them that their women worked and fought as hard as their men, and that's also true.
https://en.wikipedia.org/wiki/Ligures

In terms of the Ligures of the modern era the hypothesis has also been that there were migrations from Gallic speaking populations in the historical era, which is why they were sometimes referred to as Celt-Ligurians.
 
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@Boreas,
I'd say the European Neolithic farmer, with a chunk of WHG picked up in Europe. I do think something approaching it has been labeled "Mediterranean" in some admixture runs, but in my opinion if you want to know how much of some ancient group is in you the comparison should be with the ancient genome.

For example, there's more of that "cluster" in southern Europeans, including Iberians who border the Atlantic as well as the Mediterranean, than there is in Turks and Levantines who also border the Mediterranean.

Take a look at this Gamba et al graphic. Look at the overlap between the Sardinians and not only the Early Neolithic Europeans like the Hungarian Neolithic sample, but Otzi and even Copper Age central Europe. This is why I think it may not be accurate to imagine this as a situation where there were hunter-gatherers on the island and then Neolithic people from the mainland arrived. The authors themselves argue against assuming this is correct. It could have been an already admixed group from the mainland, perhaps the French coast, but also, as I once argued, perhaps from the southeast, and even later. I don't think this subclade of I2a1 is particularly "old" is it? On the other hand, there's the Basque affinity to consider. I wish they had done more comparison with samples like CO1.*

In comparing this study with prior ones we've all looked at for a long time, we have to keep in mind that these samples are not the ones chosen by Cavalli-Sforza and used subsequently. The authors got verbal information from Piazza as to where those were taken, and although a lot of them were from this isolated region there were some from peripheral groups as well. There's a lot of overlap, but they're not precisely the same. I'd like to see some of the studies redone with these Massif samples to see if there are any changes at all.

ncomms6257-f2.jpg
 
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I just saw these on anthrogenica on the thread for the new Sardinian paper. It was posted by Gravetto Danubian. I like most of his posts a lot. I hope he doesn’t mind my reproducing them here.

French: (It would be nice to see French Southwest)

French_East
"Bell_Beaker_Czech" 39.35
"Armenia_EBA" 24.25
"Baalberge_MN" 17.65
"Hungary_HG" 7.5
"Jordan_EBA" 6.1

Italian_Tuscan (It would be interesting to see the other Italians as well.)
"Bell_Beaker_Czech" 31.3
"Armenia_EBA" 29.05
"Baalberge_MN" 17.85
"LBK_EN" 13.7
"Jordan_EBA" 5.35

Spanish (I think this is the half Catalan sample.)
"Bell_Beaker_Czech" 37.95
"Armenia_EBA" 17.9
"Baalberge_MN" 14.35
"Jordan_EBA" 10.45
"Israel_Natufian" 5.55


Now take a look at Basques and Sardinians
Basque_French
"Bell_Beaker_Czech" 38.4
"Baalberge_MN" 26.85
"Remedello_BA" 17.7
"Armenia_EBA" 6.3
"LaBrana1" 5.6

Sardinian
"Baalberge_MN" 28
"Remedello_BA" 25.3
"Armenia_EBA" 15.9
"Anatolia_Neolithic" 10.5
"Iberia_Chalcolithic" 10
"Jordan_EBA" 6.7

(I’m assuming this is the standard Sardinian sample, not the one used in the Chiang Sardinian paper. The standard one includes some samples outside of the isolated massif samples, although they’re not the majority.)

My take away is that given the Czech Bell Beaker numbers, which is a stand in the group moving into Europe with steppe ancestry (although it’s not perfect), the Sardinians don’t have any. Everyone else in Europe has a “West Asian” component, but the Basques have very little of it.

So, the question is when did it arrive and with whom" One clue is that it’s at 19% in the British and 29% in Tuscans. So, could we conclude that at least around 19% must have been in Europe for a long time? I'm aware that these clusters are different from run to run, but dodecad found "West Asian" already in Otzi in the Copper Age, so perhaps there were slightly different migrations in the late Copper Age. Assuming that's correct, where did the other 10 points come from and when? We really need more ancient dna from Italy, Greece, the Aegean etc.

Oh, this is the result for Kent in England, which has the most Bell Beaker.
English_Kent
"Bell_Beaker_Czech" 55.95
"Armenia_EBA" 19.2
"Baalberge_MN" 13.85
"Hungary_HG" 4.25
"LaBrana1" 4.05
 

I've missed this post, sorry

regarding Mesolithic : http://eprints.uniss.it/6328/1/Melis_P_Approdo_della_costa_di.pdf

Ad epoca preneolitica, circa 9.000 anni fa (data calibrata), dovrebbe datarsi anche un primo episodio di colonizzazione dell'Isola ad opera di genti arrivate in Corsica e Sardegna dopo un'impegnativa navigazione, portatrici di caratteri culturali mesolitici ascrivibili al cosiddetto epipaleolitico indifferenziato della penisola italiana e recentemente riconosciuti nella zona di Perfugas e Laerru, nella Sardegna settentrionale.

Another paper about Mesolithic Sardinia and Corsica is Fabio Martini, Carlo Tozzi, Il mesolitico in Sardegna (there is a PDF online)

sardegnacorsica.png


Neolithic:https://web.archive.org/web/2013092...Dispense e tavole scaricabili/neolitico_4.pdf

Notevoli affinità con il repertorio formale della Sardegna e della Corsica (stile di Basi) hanno originato la definizione di stile di “Basi-Pienza” e inducono a ipotizzare una colonizzazione di queste isole maggiori da parte di comunità stanziate nell’Italia tirrenica, attraverso l’Arcipelago Toscano. L’esistenza di contatti è, del resto, confermata dalla presenza di ossidiana sarda dal Monte Arci negli assemblage litici della ceramica impressa tirrenica.

apparently a new paper with aDNA from Sardinia will be published soon
https://ep70.eventpilotadmin.com/web/page.php?page=IntHtml&project=ASHG17&id=170122074

From Neolithic to Bronze Age they found relative stability, so probably they are talking about EEF with maybe a bit of steppe admxiture in Copper/Bronze Age. (?) :unsure:
 
I've missed this post, sorry

regarding Mesolithic : http://eprints.uniss.it/6328/1/Melis_P_Approdo_della_costa_di.pdf

Ad epoca preneolitica, circa 9.000 anni fa (data calibrata), dovrebbe datarsi anche un primo episodio di colonizzazione dell'Isola ad opera di genti arrivate in Corsica e Sardegna dopo un'impegnativa navigazione, portatrici di caratteri culturali mesolitici ascrivibili al cosiddetto epipaleolitico indifferenziato della penisola italiana e recentemente riconosciuti nella zona di Perfugas e Laerru, nella Sardegna settentrionale.

Another paper about Mesolithic Sardinia and Corsica is Fabio Martini, Carlo Tozzi, Il mesolitico in Sardegna (there is a PDF online)

View attachment 9534


Neolithic:https://web.archive.org/web/2013092...Dispense e tavole scaricabili/neolitico_4.pdf

Notevoli affinità con il repertorio formale della Sardegna e della Corsica (stile di Basi) hanno originato la definizione di stile di “Basi-Pienza” e inducono a ipotizzare una colonizzazione di queste isole maggiori da parte di comunità stanziate nell’Italia tirrenica, attraverso l’Arcipelago Toscano. L’esistenza di contatti è, del resto, confermata dalla presenza di ossidiana sarda dal Monte Arci negli assemblage litici della ceramica impressa tirrenica.

apparently a new paper with aDNA from Sardinia will be published soon
https://ep70.eventpilotadmin.com/web/page.php?page=IntHtml&project=ASHG17&id=170122074


Grazie. Yes, the obsidian tells us a lot. I had a long running argument with Jean Manco about the old sites on Sardinia. She always maintained they were only temporary sites, not settlements. Maybe, maybe not.
 
452px-Obsidian_diffusion_Mediterranean_map.svg.png

but the dating is a bit tricky
somewhere in the EN

it is not the mesolithic as Sardegna was virtualy uninhabited before Cardial Ware

only few scarce settlements that were probably inhabited not more than a few generations maximum

https://www.nature.com/articles/srep42869

The earliest presence of humans is still under debate. Some authors likely date it back to the end of the Middle Pleistocene, on the base of lithic artifacts typology, attributed to the Lower Paleolithic20,21,22. Nonetheless, neither human remains nor absolute-dated contextual evidence support this hypothesis. However, clues of human settlements arose only from the end of the Upper Pleistocene23,24, with single human remains discovered out of context and dated back to 20,000 years ago just on the base of stratigraphic correlations25. The first evidence of Holocene frequentation of the island are scattered in a few rock-shelters and caves, exclusively on the inside of a 20 km coast belt26. After this poorly-documented phase, with around 500 years hiatus of archaeological evidence, with the advent of the agriculture, the population of the island increased in size.
 
it is not the mesolithic as Sardegna was virtualy uninhabited before Cardial Ware
only few scarce settlements that were probably inhabited not more than a few generations maximum

hmm i don't know, some Sardianian population like Gairo are +20% circa extra WHG compared to EEF

chiangsupp.png
 
not sure if already posted but this is an entire book on pre-history sardinia/(mediterranean)
http://members.peak.org/~obsidian/pdf/tykot_1995.pdf

concerning the genetics, can it be said that modern sardinians are more close(shared drift) towards 'farmers' of the kum6<ötzi/remedello type than to EN/Cardial CB13/TOR types? meaning a superseding by an add 4th mil wave from anatolia(<italy?<balkans?) as described in kilinc et al, a new wave of metallurgical skilled farmers which coincide with ozieri and the late5th/4th mil emerging copper production/network and slim-mother idols of an akin cyclade/anatolian type (in contrast to previous fat-mother bonu ighinu); though from what i have seen the sardinian copper network was most active during the 2nd mil (~1500bc) bronze age as the main source for all of italy and a source/part of a network as far as scandinavia;
 
^^Jean Manco speculated long ago on dna forums that the majority of Sardinia's genetic make up stems from a migration from the Balkans of metallurgists. I agreed with her then and I still think it's a decent option.

The reason that they look so much like the Early Farmers genetically is that the people in the Balkans during the Copper Age probably weren't that different from the Early Farmers. It would explain why Otzi plots with them, and Copper Age Baden isn't so very different either.
 
could have also been a direct migration from anatolia (copper driven) since the CBMP was east/steppe-belt oriented and collapsed during the late 5th early 4th; kumtepe itself is a site of copper objects and more important processing and deposit[troas] but it is not solely metallurgical based, per aDNA the new wave is marked by the shared alleles/drift with CHG driven westward by tepecik (not present in previous bonckulu or contemporary barcin) > kumtepe and known >italy (ötzi/remedello) who belonged to a copper network (production/innovation) of tuscany; balkan_Chl in qpAdm mathieson et al do have ~6%CHG and the VBQ change is marked by an extensive trans-alpine trade (adige<eisack<inn) where characteristics like crouched (single-grave) burials and type-arrowheads are also later part of the remedello horizon; in toto either a direct metallurgical/copper driven migration antolia>sardinia (pos sardinia>tuscany) or via balkan>italy/(lunigiana)>sardinia but maybe even a mash up of carpathian_farmers and anatolian_metallurgists (via <tuscany<sardinia) in north_italy at least; a VBQ burials
http://www.academia.edu/1456884/Tes...milia_Occidentale._Archeologia_e_antropologia
12qyLHX.png
 

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