Ancient Sardinia hints at Mesolithic spread of R1b-V88, and Western EEF-related expansion of Vasconic

nuragic-sardinia-neolithic

New preprint Population history from the Neolithic to present on the Mediterranean island of Sardinia: An ancient DNA perspective, by Marcus et al. bioRxiv (2019)

Interesting excerpts (emphasis mine, edited for clarity):

On the high frequency of R1b-V88

Our genome-wide data allowed us to assign Y haplogroups for 25 ancient Sardinian individuals. More than half of them consist of R1b-V88 (n=10) or I2-M223 (n=7).

Francalacci et al. (2013) identi fied three major Sardinia-specifi c founder clades based on present-day variation within the haplogroups I2-M26, G2-L91 and R1b-V88, and here we found each of those broader haplogroups in at least one ancient Sardinian individual. Two major present-day Sardinian haplogroups, R1b-M269 and E-M215, are absent.

Compared to other Neolithic and present-day European populations, the number of identi fied R1b-V88 carriers is relatively high.

(…)ancient Sardinian mtDNA haplotypes belong almost exclusively to macro-haplogroups HV (n = 16), JT (n = 17) and U (n = 9), a composition broadly similar to other European Neolithic populations.

r1b-v88-europe
Geographic and temporal distribution of R1b-V88 Y-haplotypes in ancient European samples. We plot the geographic position of all ancient samples inferred to carry R1b-V88 equivalent markers. Dates are given as years BCE (means of calibrated 2s radio-carbon dates). Multiple V88 individuals with similar geographic positions are vertically stacked. We additionally color-code the status of the R1b-V88 subclade R1b-V2197, which is found in most present-day African R1b-V88 carriers.

On the origin of a Vasconic-like Paleosardo with the Western EEF

(…) the Neolithic (and also later) ancient Sardinian individuals sit between early Neolithic Iberian and later Copper Age Iberian populations, roughly on an axis that differentiates WHG and EEF populations and embedded in a cluster that additionally includes Neolithic British individuals. This result is also evident in terms of absolute genetic differentiation, with low pairwise FST ~ 0.005 +- 0.002 between Neolithic Sardinian individuals and Neolithic western mainland European populations. Pairwise outgroup-f3 analysis shows a very similar pattern, with the highest values of f3 (i.e. most shared drift) being with Neolithic and Copper Age Iberia, gradually dropping off for temporally and geographically distant populations.

In explicit admixture models (using qpAdm, see Methods) the southern French Neolithic individuals (France-N) are the most consistent with being a single source for Neolithic Sardinia (p ~ 0:074 to reject the model of one population being the direct source of the other); followed by other populations associated with the western Mediterranean Neolithic Cardial Ware expansion.

sardinians-ancient-eef
Principal Components Analysis based on the Human Origins dataset. A: Projection of ancient individuals’ genotypes onto principal component axes de fined by modern Western Eurasians (gray labels).

Pervasive Western Hunter-Gatherer ancestry in Iberian/French/Sardinian population

Similar to western European Neolithic and central European Late Neolithic populations, ancient Sardinian individuals are shifted towards WHG individuals in the top two PCs relative to early Neolithic Anatolians Admixture analysis using qpAdm infers that ancient Sardinian individuals harbour HG ancestry (~ 17%) that is higher than early Neolithic mainland populations (including Iberia, ~ 8%), but lower than Copper Age Iberians (~ 25%) and about the same as Southern French Middle-Neolithic individuals (~ 21%).

sardinia-modern-ancient-nuragic-pca
Principal Components Analysis based on the Human Origins dataset. B: Zoom into the region most relevant for Sardinian individuals.

Continuity from Sardinia Neolithic through the Nuragic

We found several lines of evidence supporting genetic continuity from the Sardinian Neolithic into the Bronze Age and Nuragic times. Importantly, we observed low genetic differentiation between ancient Sardinian individuals from various time periods.

A qpAdm analysis, which is based on simultaneously testing f-statistics with a number of outgroups and adjusts for correlations, cannot reject a model of Neolithic Sardinian individuals being a direct predecessor of Nuragic Sardinian individuals (…) Our qpAdm analysis further shows that the WHG ancestry proportion, in a model of admixture with Neolithic Anatolia, remains stable at ~17% throughout three ancient time-periods.

sardinians-modern-ancient-pca-admixture
Present-day genetic structure in Sardinia reanalyzed with aDNA. A: Scatter plot of the rst two principal components trained on 1577 present-day individuals with grand-parental ancestry from Sardinia. Each individual is labeled with a location if at least 3 of the 4 grandparents were born in the same geographical location (\small” three letter abbreviations); otherwise with \x” or if grand-parental ancestry is missing with \?”. We calculated median PC values for each Sardinian province (large abbreviations). We also projected each ancient Sardinian individual on to the top two PCs (gray points). B/C: We plot f-statistics that test for admixture of modern Sardinian individuals (grouped into provinces) when using Nuragic Sardinian individuals as one source population. Uncertainty ranges depict one standard error (calculated from block bootstrap). Karitiana are used in the f-statistic calculation as a proxy for ANE/Steppe ancestry (Patterson et al., 2012).

Steppe influx in Modern Sardinians

While contemporary Sardinian individuals show the highest affinity towards EEF-associated populations among all of the modern populations, they also display membership with other clusters (Fig. 5). In contrast to ancient Sardinian individuals, present-day Sardinian individuals carry a modest “Steppe-like” ancestry component (but generally less than continental present-day European populations), and an appreciable broadly “eastern Mediterranean” ancestry component (also inferred at a high fraction in other present-day Mediterranean populations, such as Sicily and Greece).

Related

Arrival of steppe ancestry with R1b-P312 in the Mediterranean: Balearic Islands, Sicily, and Iron Age Sardinia

steppe-balearic-sicily-sardinia

New preprint The Arrival of Steppe and Iranian Related Ancestry in the Islands of the Western Mediterranean by Fernandes, Mittnik, Olalde et al. bioRxiv (2019)

Interesting excerpts (emphasis in bold; modified for clarity):

Balearic Islands: The expansion of Iberian speakers

Mallorca_EBA dates to the earliest period of permanent occupation of the islands at around 2400 BCE. We parsimoniously modeled Mallorca_EBA as deriving 36.9 ± 4.2% of her ancestry from a source related to Yamnaya_Samara; (…). We next used qpAdm to identify “proximal” sources for Mallorca_EBA’s ancestry that are more closely related to this individual in space and time, and found that she can be modeled as a clade with the (small) subset of Iberian Bell Beaker culture associated individuals who carried Steppe-derived ancestry (p=0.442).

Suppl. Materials: The model used was with Bell_Beaker_Iberia_highsteppe, a group of outliers from Iberia buried in a Bell Beaker mortuary context who unlike most individuals from this context in that region had high proportions of Steppe ancestry (p=0.442).

Our estimates of Steppe ancestry in the two later Balearic Islands individuals are lower than the earlier one: 26.3 ± 5.1% for Formentera_MBA and 23.1 ± 3.6% for Menorca_LBA, but the Middle to Late Bronze Age Balearic individuals are not a clade relative to non-Balearic groups. Specifically, we find that f4(Mbuti.DG, X; Formentera_MBA, Menorca_LBA) is positive when X=Iberia_Chalcolithic (Z=2.6) or X=Sardinia_Nuragic_BA (Z=2.7). While it is tempting to interpret the latter statistic as suggesting a genetic link between peoples of the Talaiotic culture of the Balearic islands and the Nuragic culture of Sardinia, the attraction to Iberia_Chalcolithic is just as strong, and the mitochondrial haplogroup U5b1+16189+@16192 in Menorca_LBA is not observed in Sardinia_Nuragic_BA but is observed in multiple Iberia_Chalcolithic individuals. A possible explanation is that both the ancestors of Nuragic Sardinians and the ancestors of Talaiotic people from the Balearic Islands received gene flow from an unsampled Iberian Chalcolithic-related group (perhaps a mainland group affiliated to both) that did not contribute to Formentera_MBA.

This sample, like another one in El Argar, is of hg. R1b-P312. So there you are, the data that connects the Proto-Iberian expansion (replacing IE-speaking Bell Beakers) to the Iberian Chalcolithic population, signaled by the increase in Iberian Chalcolithic ancestry after the arrival of Bell Beakers, most likely connected originally to the Argaric and post-Argaric expansions during the MBA.

balearic-sicily-sardinia-pca
PCA with previously published ancient individuals (non-filled symbols), projected onto variation from present-day populations (gray squares).

Steppe in Sardinia IA: Phocaeans from Italy?

Most Sardinians buried in a Nuragic Bronze Age context possessed uniparental haplogroups found in European hunter-gatherers and early farmers, including Y-haplogroup R1b1a[xR1b1a1a] which is different from the characteristic R1b1a1a2a1a2 spread in association with the Bell Beaker complex. An exception is individual I10553 (1226-1056 calBCE) who carried Y-haplogroup J2b2a, previously observed in a Croatian Middle Bronze Age individual bearing Steppe ancestry, suggesting the possibility of genetic input from groups that arrived from the east after the spread of first farmers. This is consistent with the evidence of material culture exchange between Sardinians and mainland Mediterranean groups, although genome-wide analyses find no significant evidence of Steppe ancestry so the quantitative demographic impact was minimal.

Another interesting data, these (Mesolithic) remnant R1b-V88 lineages closely related to the Italian Peninsula, the most likely region of expansion of these lineages into Africa, in turn possibly connected to the expansion of Proto-Afroasiatic.

We detect definitive evidence of Iranian-related ancestry in an Iron Age Sardinian I10366 (391-209 calBCE) with an estimate of 11.9 ± 3.7.% Iran_Ganj_Dareh_Neolithic related ancestry, while rejecting the model with only Anatolian_Neolithic and WHG at p=0.0066 (Supplementary Table 9). The only model that we can fit for this individual using a pair of populations that are closer in time is as a mixture of Iberia_Chalcolithic (11.9 ± 3.2%) and Mycenaean (88.1 ± 3.2%) (p=0.067). This model fits even when including Nuragic Sardinians in the outgroups of the qpAdm analysis, which is consistent with the hypothesis that this individual had little if any ancestry from earlier Sardinians.

yamnaya-samara
Proportions of ancestry using a distal qpAdm framework on an individual basis (a), and based on qpWave clusters

Sicily EBA: The Lusitanian/Ligurian connection?

(…) While a previously reported Bell Beaker culture-associated individual from Sicily had no evidence of Steppe ancestry, (…) we find evidence of Steppe ancestry in the Early Bronze Age by ~2200 BCE. In distal qpAdm, the outlier Sicily_EBA11443 is parsimoniously modeled as harboring 40.2 ± 3.5% Steppe ancestry, and the outlier Sicily_EBA8561 is parsimoniously modeled as harboring 23.3 ± 3.5% Steppe ancestry. (…) The presence of Steppe ancestry in Early Bronze Age Sicily is also evident in Y chromosome analysis, which reveals that 4 of the 5 Early Bronze Age males had Steppe-associated Y-haplogroup R1b1a1a2a1a2. (Online Table 1). Two of these were Y-haplogroup R1b1a1a2a1a2a1 (Z195) which today is largely restricted to Iberia and has been hypothesized to have originated there 2500-2000 BCE. This evidence of west-to-east gene flow from Iberia is also suggested by qpAdm modeling where the only parsimonious proximate source for the Steppe ancestry we found in the main Sicily_EBA cluster is Iberians.

What’s this? An ancestral connection between Sicel Elymian and Galaico-Lusitanian or Ligurian (based on an origin in NE Iberia)? Impossible to say, especially if the languages of these early settlers were replaced later by non-Indo-European speakers from the eastern Mediterranean, and by Indo-European speakers from the mainland closely related to Proto-Italic during the LBA, but see below.

Regarding the comment on R1b-Z195, it is associated with modern Iberians, as DF27 in general, due to founder effects beyond the Pyrenees. It is a very old subclade, split directly from DF27 roughly at the same time as it split from the parent P312, i.e. it can be found anywhere in Europe, and it almost certainly accompanied the expansion of Celts from Central Europe under the subclade R1b-M167/SRY2627.

The connection is thus strong only because of the qpAdm modeling, since R1b-DF27 and subclade R1b-Z195 are certainly lineages expanded quite early, most likely with Yamna settlers in Hungary and East Bell Beakers.

In this case, if stemming from Iberia, it is most likely of subclade R1b-Z220 – or another Z195 (xM167) lineage – originally associated with the Old European substrate found in topo-hydronymy in Iberia, whose most likely remnants attested during the Iron Age were Lusitanians.

r1b-df27-z195
Left: Modern distribution of R1b-Z195 (YFull estimate 2700 BC); Right: Modern distribution of DF27. Both include later founder effects within Iberia, so the increase in the Basque country and the Crown of Aragon and the decrease in Portugal can safely be ignored. Contour maps of the derived allele frequencies of the SNPs analyzed in Solé-Morata et al. (2017).

We detect Iranian-related ancestry in Sicily by the Middle Bronze Age 1800-1500 BCE, consistent with the directional shift of these individuals toward Mycenaeans in PCA. Specifically, two of the Middle Bronze Age individuals can only be fit with models that in addition to Anatolia_Neolithic and WHG, include Iran_Ganj_Dareh_Neolithic. The most parsimonious model for Sicily_MBA3125 has 18.0 ± 3.6% Iranian-related ancestry (p=0.032 for rejecting the alternative model of Steppe rather than Iranian-related ancestry), and the most parsimonious model for Sicily_MBA has 14.9 ± 3.9% Iranian-related ancestry (p=0.037 for rejecting the alternative model).

The modern southern Italian Caucasus-related signal identified in Raveane et al. (2018) is plausibly related to the same Iranian-related spread of ancestry into Sicily that we observe in the Middle Bronze Age (and possibly the Early Bronze Age).

The non-Indo-European Sicanians and Elymians were possibly then connected to eastern Mediterranean groups before the expansion of the Sea Peoples.

For the Late Bronze Age group of individuals, qpAdm documented Steppe-related ancestry, modeling this group as 80.2 ± 1.8% Anatolia_Neolithic, 5.3 ± 1.6% WHG, and 14.5 ± 2.2% Yamnaya_Samara. Our modeling using sources more closely related in space and time also supports Sicily_LBA having Minoan-related ancestry or being derived from local preceding populations or individuals with ancestries similar to those of Sicily_EBA3123 (p=0.527), Sicily_MBA3124 (p=0.352), and Sicily_MBA3125 (p=0.095).

This increase in Steppe-related ancestry in a western site during the LBA most likely represents either an expansion from the Aegean or – maybe more likely, given the archaeological finds – a regional population similar to Sicily EBA re-emerging or rather being displaced from the eastern part of the island because of a westward movement from nearby Calabria.

Whether this population sampled spoke Indo-European or not at this time is questionable, since the Iron Age accounts show non-IE Elymians in this region.

Actually, Elymians seem to have spoken Indo-European, which fits well with the increase in steppe ancestry.

EDIT (21 MAR): Interesting about a proposed incoming Minoan-like ancestry is the potential origin of the Iran Neolithic-related ancestry that is going to appear in Central Italy during the LBA. This could then be potentially associated with Tyrsenians passing through the area, although the traditional description may be more more compatible with an arrival of Sea Peoples from the Adriatic.

Sad to read this:

This manuscript is dedicated to the memory of Sebastiano Tusa of the Soprintendenza del Mare in Palermo, who would have been an author of this study had he not tragically died in the crash of Ethiopia Airlines flight 302 on March 10.

Related

Happy new year 2019…and enjoy our new books!

song-sheep-horses-header

Sorry for the last weeks of silence, I have been rather busy lately. I am having more projects going on, and (because of that) I also wanted to finish a project I have been working on for many months already.

I have therefore decided to publish a provisional version of the text, in the hope that it will be useful in the following months, when I won’t be able to update it as often as I would like to:

EDIT (20 JAN 2019): For those of you who are more comfortable reading in your native language, I have placed some links to automatic translations by Google Translate. They might work especially well for the texts of A Game of Clans & A Clash of Chiefs.

Don’t forget to check out the maps included in the supplementary materials: I have added Y-DNA, mtDNA, and ADMIXTURE data using GIS software. The PCA graphics are also important to follow the main text.

NOTE. Right now the files are only in my server. I will try to upload them to Academia.edu and Research Gate when I have time, I have uploaded them to Academia.edu and ResearchGate, in case the websites are too slow.

I would have preferred to wait for a thorough revision of the section on archaeology and the linguistic sections on Uralic, but I doubt I will have time when the reviews come, so it was either now or maybe next December…

I say so in the introduction, but it is evident that certain aspects of the book are tentative to say the least: the farther back we go from Late Proto-Indo-European, the less clear are many aspects. Also, linguistically I am not convinced about Eurasiatic or Nostratic, although they do have a certain interest when we try to offer a comprehensive view of the past, including ethnolinguistic identities.

I cannot be an expert in everything, and these books cover a lot. I am bound to publish many corrections as new information appears and more reviews are sent. For example, just days ago (before SNP calls of Wang et al. 2018 were published) some paragraphs implied that AME might have expanded Nostratic from the Middle East. Now it does not seem so, and I changed them just before uploading the text. That’s how tentative certain routes are, and how much all of this may change. And that only if we accept a Nostratic phylum…

NOTE. Since the first book I wrote was the linguistic one, and I have spent the last months updating the archaeology + genetics part, now many of you will probably understand 1) why I am so convinced about certain language relationships and 2) how I used many posts to clarify certain ideas and receive comments. Many posts offer probably a good timeline of what I worked with, and when.

Acknowledgements

I did not add this section to the books, because they are still not ready for print, but I think this is due somewhere now. It is impossible to reference all who have directly or indirectly contributed to this, so this is a list of those I feel have played an important role.

I am indebted to the following people (which does not mean that they share my views, obviously):

First and foremost, to Fernando López-Menchero, for having the patience to review with detail many parts on Indo-European linguistics, knowing that I won’t accept many of his comments anyway. The additional information he offers is invaluable, but I didn’t want to turn this into a huge linguistic encyclopaedia with unending discussions of tiny details of each reconstructed word. I think it is already too big as it is.

I would not have thought about doing this if it were not for the interest of Wekwos (Xavier Delamarre) in publishing a full book about the Indo-European demic diffusion model (in the second half of 2017, I think). It was them who suggested that I extended the content, when all I had done until then was write an essay and draw some maps in my free time between depositing the PhD thesis and defending it.

Sadly, as much as I would like to publish a book with a professional publisher, I don’t think ancient DNA lends itself for the traditional format, so my requests (mainly to have free licenses and being able to review the text at will, as new genetic papers are published) were logically not acceptable. Also, the main aim of all volumes, especially the linguistic one, is the teaching of essentials of Late Proto-Indo-European and related languages, and this objective would be thwarted by selling each volume for $50-70 and only in printed format. I prefer a wider distribution.

At first I didn’t think much of this proposal, because I do not benefit from this kind of publications in my scientific field, but with time my interest in writing a whole, comprehensive book on the subject grew to the point where it was already an ongoing project, probably by the start of 2018.

I would not have been in contact with Wekwos if it were not for user Camulogène Rix at Anthrogenica, so thanks for that and for the interest in this work.

I would not have thought of writing this either if not for the spontaneous support (with an unexpected phone call!) of a professor of the Complutense University of Madrid, Ángel Gómez Moreno, who is interested in this subject – as is his wife, a professor of Classics more closely associated to Indo-European studies, and who helped me with a search for Indo-Europeanists.

EDIT (1 JAN 2019): I remembered that Karin Bojs sent me her book after reading the demic diffusion model. I may have also thought about writing a whole book back then, but mid-2017 is probably too early for the project.

Professor Kortlandt is still to review the text, but he contributed to both previous essays in some very interesting ways, so I hope he can help me improve the parts on Uralic, and maybe alternative accounts of expansion for Balto-Slavic, depending on the time depth that he would consider warranted according to the Temematic hypothesis.

The maps are evidently (for those who are interested in genetics) in part the result of the effort of the late Jean Manco: As you can see from the maps including Y-DNA and mtDNA samples, I have benefitted from her way of organising data and publishing it. Similarly, the work of Iain McDonald in assessing the potential migration routes of R1b and R1a in Europe with the help of detailed maps was behind my idea for the first maps, and consequently behind these, too.

I should thank all people responsible for the release of free datasets to work with, including the Reich and Jena labs, the Veeramah Lab, and also researchers from the Max Planck Institute or the Mainz Palaeogenetics group, who didn’t mind to share with me datasets to work with.

Readers of this blog with interesting comments have also been essential for the improvement of the texts. You can probably see some of your many contributions there. I may not answer many comments, because I am always busy (and sometimes I just don’t have anything interesting to say), but I try to read all of them.

EDIT (1 JAN 2019) I think I should mention at least Chetan, Egg, or Robert George; but then I would leave out old europe, Sgr Ganesh, or Tileman Ehlen; and if I include them I would leave out others…

Users of other sites, like Anthrogenica, whose particular points of view and deep knowledge of some very specific aspects are sometimes very useful. In particular, user Anglesqueville helped me to fix some issues with the merging of datasets to obtain the PCAs and ADMIXTURE, and prepared some individual samples to merge them.

Even without posting anything, Google Analytics keeps sending me messages about increasing user fidelity (returning users), and stats haven’t really changed (which probably means more people are reading old posts), so thank you for that.

I hope you enjoy the books.

Happy new year!

R1a-Z280 lineages in Srubna; and first Palaeo-Balkan R1b-Z2103?

herodotus-world-map

Scythian samples from the North Pontic area are far more complex than what could be seen at first glance. From the new Y-SNP calls we have now thanks to the publications at Molgen (see the spreadsheet) and in Anthrogenica threads, I think this is the basis to work with:

NOTE. I understand that writing a paper requires a lot of work, and probably statistical methods are the main interest of authors, editors, and reviewers. But it is difficult to comprehend how any user of open source tools can instantly offer a more complex assessment of the samples’ Y-SNP calls than professionals working on these samples for months. I think that, by now, it should be clear to everyone that Y-DNA is often as important (sometimes even more) than statistical tools to infer certain population movements, since admixture can change within few generations of male-biased migrations, whereas haplogroups can’t…

Srubna

Srubna-Andronovo samples are as homogeneous as they always were, dominated by R1a-Z645 subclades and CWC-related (steppe_MLBA) ancestry.

The appearance of one (possibly two) R-Z280 lineages in this mixed Srubna-Alakul region of the southern Urals and this early (1880-1690 BC, hence rather Pokrovka-Alakul) points to the admixture of R1a-Z93 and R1a-Z280 already in Abashevo, which also explains the wide distribution of both subclades in the forest zones of Central Asia.

If Abashevo is the cornerstone of the Indo-Iranian / Uralic community, as it seems, the genetic admixture would initially be quite similar, undergoing in the steppes a reduction to haplogroup R1a-Z93 (obviously not complete), at the same time as it expanded to the west with Pokrovka and Srubna, and to the east with Petrovka and Andronovo. To the north, similar reductions will probably be seen following the Seima-Turbino phenomenon.

NOTE. Another R1a-Z280 has been found in the recent sample from Bronze Age Poland (see spreadsheet). As it appears right now in ancient and modern DNA, there seems to be a different distribution between subclades:

  • R1a-Z280 (formed ca. 2900 BC, TMRCA ca. 2600 BC) appears mainly distributed today to the east, in the forest and steppe regions, with the most ‘successful’ expansions possibly related to the spread of Abashevo- and Battle Axe-related cultures (Indo-Iranian and Uralic alike).
  • R1a-M458 (formed ca. 2700, TMRCA ca. 2700 BC) appears mainly distributed to the north, from central Europe to the east – but not in the steppe in aDNA, with the most ‘successful’ expansions to the west.

M458 lineages seem thus to have expanded in the steppe in sizeable numbers only after the Iranian expansions (see a map of modern R1a distributions) i.e. possibly with the expansion of Slavs, which supports the model whereby cultures from central-east Europe (like Trzciniec and Lusatian), accompanied mainly by M458 lineages, were responsible for the expansion of Proto-Balto-Slavic (and later Proto-Slavic).

The finding of haplogroup R1a-Z93, among them one Z2123, is no surprise at this point after other similar Srubna samples. As I said, the early Srubna expansion is most likely responsible for the Szólád Bronze Age sample (ca. 2100-1700 BC), and for the Balkans BA sample (ca. 1750-1625 BC) from Merichleri, due to incursions along the central-east European steppe.

cheek-pieces
Map of decorated bone/antler bridle cheek-pieces and whip handle equivalents. They are often local translations that remained faithful to the originals (from data in Piggott, 1965; Kristiansen & Larsson, 2005; David, 2007). Image from Vandkilde (2014).

Cimmerians

Cimmerian samples from the west show signs of continuity with R1a-Z93 lineages. Nevertheless, the sample of haplogroup Q1a-Y558, together with the ‘Pre-Scythian’ sample of haplogroup N (of the Mezőcsát Culture) in Hungary ca. 980-830 BC, as well as their PCA, seem to depict an origin of these Pre-Scythian peoples in populations related to the eastern Central Asian steppes, too.

NOTE. I will write more on different movements (unrelated to Uralic expansions) from Central and East Asia to the west accompanied by Siberian ancestry and haplogroup N with the post of Ugric-Samoyedic expansions.

Scythians

The Scythian of Z2123 lineage ca. 375-203 BC from the Volga (in Mathieson et al. 2015), together with the sample scy193 from Glinoe (probably also R1a-Z2123), without a date, as well as their common Steppe_MLBA cluster, suggest that Scythians, too, were at first probably quite homogeneous as is common among pastoralist nomads, and came thus from the Central Asian steppes.

The reduction in haplogroup variability among East Iranian peoples seems supported by the three new Late Sarmatian samples of haplogroup R1a-Z2124.

Approximate location of Glinoe and Glinoe Sad (with Starosilya to the south, in Ukrainian territory):

This initial expansion of Scythians does not mean that one can dismiss the western samples as non-Scythians, though, because ‘Scythian’ is a cultural attribution, based on materials. Confirming the diversity among western Scythians, a session at the recent ISBA 8:

Genetic continuity in the western Eurasian Steppe broken not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths), by Järve et al.

The long-held archaeological view sees the Early Iron Age nomadic Scythians expanding west from their Altai region homeland across the Eurasian Steppe until they reached the Ponto-Caspian region north of the Black and Caspian Seas by around 2,900 BP. However, the migration theory has not found support from ancient DNA evidence, and it is still unclear how much of the Scythian dominance in the Eurasian Steppe was due to movements of people and how much reflected cultural diffusion and elite dominance. We present new whole-genome results of 31 ancient Western and Eastern Scythians as well as samples pre- and postdating them that allow us to set the Scythians in a temporal context by comparing the Western Scythians to samples before and after within the Ponto-Caspian region. We detect no significant contribution of the Scythians to the Early Iron Age Ponto-Caspian gene pool, inferring instead a genetic continuity in the western Eurasian Steppe that persisted from at least 4,800–4,400 cal BP to 2,700–2,100 cal BP (based on our radiocarbon dated samples), i.e. from the Yamnaya through the Scythian period.

(…) Our results (…) support the hypothesis that the Scythian dominance was cultural rather than achieved through population replacement.

Detail of the slide with admixture of Scythian groups in Ukraine:

scythians-admixture

The findings of those 31 samples seem to support what Krzewińska et al. (2018) found in a tiny region of Moldavia-south-western Ukraine (Glinoi, Glinoi Sad, and Starosilya).

The question, then, is as follows: if Scythian dominance was “cultural rather than achieved through population replacement”…Where are the R1b-Z2103 from? One possibility, as I said in the previous post, is that they represent pockets of Iranian R1b lineages in the steppes descended from eastern Yamna, given that this haplogroup appears in modern populations from a wide region surrounding the steppes.

The other possibility, which is what some have proposed since the publication of the paper, is that they are related to Thracians, and thus to Palaeo-Balkan populations. About the previously published Thracian individuals in Sikora et al. (2014):

thracian-samples
Geographic origin of ancient samples and ADMIXTURE results. (A) Map of Europe indicating the discovery sites for each of the ancient samples used in this study. (B) Ancestral population clusters inferred using ADMIXTURE on the HGDP dataset, for k = 6 ancestral clusters. The width of the bars of the ancient samples was increased to aid visualization. https://doi.org/10.1371/journal.pgen.1004353.g001

For the Thracian individuals from Bulgaria, no clear pattern emerges. While P192-1 still shows the highest proportion of Sardinian ancestry, K8 more resembles the HG individuals, with a high fraction of Russian ancestry.

Despite their different geographic origins, both the Swedish farmer gok4 and the Thracian P192-1 closely resemble the Iceman in their relationship with Sardinians, making it unlikely that all three individuals were recent migrants from Sardinia. Furthermore, P192-1 is an Iron Age individual from well after the arrival of the first farmers in Southeastern Europe (more than 2,000 years after the Iceman and gok4), perhaps indicating genetic continuity with the early farmers in this region. The only non-HG individual not following this pattern is K8 from Bulgaria. Interestingly, this individual was excavated from an aristocratic inhumation burial containing rich grave goods, indicating a high social standing, as opposed to the other individual, who was found in a pit.

pca-thracians

The following are excerpts from A Companion to Ancient Thrace (2015), by Valeva, Nankov, and Graninger (emphasis mine):

Thracian settlements from the 6th c. BC on:

(…) urban centers were established in northeastern Thrace, whose development was linked to the growth of road and communication networks along with related economic and distributive functions. The early establishment of markets/emporia along the Danube took place toward the middle of the first millennium BCE (Irimia 2006, 250–253; Stoyanov in press). The abundant data for intensive trade discovered at the Getic village in Satu Nou on the right bank of the Danube provides another example of an emporion that developed along the main artery of communication toward the interior of Thrace (Conovici 2000, 75–76).

Undoubtedly the most prominent manifestation of centralization processes and stratification in the settlement system of Thrace arrives with the emergence of political capitals – the leading urban centers of various Thracian political formations.

getic-thracian
Image from Volf at Vol_Vlad LiveJournal.

Their relationships with Scythians and Greeks

The Scythian presence south of the Danube must be balanced with a Thracian presence north of the river. We have observed Getae there in Alexander’s day, settled and raising grain. For Strabo the coastlands from the Danube delta north as far as the river and Greek city of Tyras were the Desert of the Getae (7.3.14), notable for its poverty and tracklessness beyond the great river. He seems to suggest also that it was here that Lysimachus was taken alive by Dromichaetes, king of the Getae, whose famous homily on poverty and imperialism only makes sense on the steppe beyond the river (7.3.8; cf. Diod. 21.12; further on Getic possessions above the Danube, Paus. 1.9 with Delev 2000, 393, who seems rather too skeptical; on poverty, cf. Ballesteros Pastor 2003). This was the kind of discourse more familiarly found among Scythians, proud and blunt in the strength of their poverty. However, as Herodotus makes clear, simple pastoralism was not the whole story as one advanced round into Scythia. For he observes the agriculture practiced north and west of Olbia. These were the lands of the Alizones and the people he calls the Scythian Ploughmen, not least to distinguish them from the Royal Scythians east of Olbia, in whose outlook, he says, these agriculturalist Scythians were their inferiors, their slaves (Hdt. 4.20). The key point here is that, as we began to see with the Getan grain-fields of Alexander’s day, there was scope for Thracian agriculturalists to maintain their lifestyles if they moved north of the Danube, the steppe notwithstanding. It is true that it is movement in the other direction that tends to catch the eye, but there are indications in the literary tradition and, especially, in the archaeological record that there was also significant movement northward from Thrace across the Danube and the Desert of the Getae beyond it.

Greek literary sources were not much concerned with Thracian migration into Scythia, but we should observe the occasional indications of that process in very different texts and contexts. At the level of myth, it is to be remembered that Amazons were regularly considered to be of Thracian ethnicity from Archaic times onward and so are often depicted in Thracian dress in Greek art (Bothmer 1957; cf. Sparkes 1997): while they are most familiar on the south coast of the Black Sea, east of Sinope, they were also located on the north coast, especially east of the Don (the ancient Tanais). Herodotus reports an origin-story of the Sauromatians there, according to which this people had been created by the union of some Scythian warriors with Amazons captured on the south coast and then washed up on the coast of Scythia (4.110). While the story is unhistorical, it is not without importance. First, it reminds us that passage north from the Danube was not the only way that Thracians, Thracian influence, and Thracian culture might find their way into Scythia. There were many more and less circuitous routes, especially by sea, that could bring Thrace into Scythia. Secondly, the myth offered some ideological basis for the Sauromatian settlement in Thrace that Strabo records, for Sauromatians might claim a Thracian origin through their Amazon forebears. Finally, rather as we saw that Heracles could bring together some of the peoples of the region, we should also observe that Ares, whose earthly home was located in Thrace by a strong Greek and Roman tradition, seems also to have been a deity of special significance and special cult among the Scythians. So much was appropriate, especially from a Classical perspective, in associations between these two peoples, whose fame resided especially in their capacity for war.

skythen
Scythians: cultures and findings (ca. 7th-4th/3rd c. BC). Greek colonies marked with concentric circles.

This broad picture of cultural contact, interaction, and osmosis, beyond simple conflict, provides the context for a range of archaeological discoveries, which – if examined separately – may seem to offer no more than a scatter of peculiarities. Here we must acknowledge especially the pioneering work of Melyukova, who has done most to develop thinking on Thracian–Scythian interaction. As she pointed out, we have a good example of Thracian–Scythian osmosis as early as the mid-seventh century bce at Tsarev Brod in northeastern Bulgaria, where a warrior’s burial combines elements of Scythian and Thracian culture (Melyukova 1965). For, while the manner of his burial and many of the grave goods find parallels in Scythia and not Thrace, there are also goods which would be odd in a Scythian burial and more at home in a Thracian one of this period (notably a Hallstatt vessel, an iron knife, and a gold diadem). Also interesting in this regard are several stone figures found in the Dobrudja which resemble very closely figures of this kind (baby) known from Scythia (Melyukova 1965, 37–38). They range in date from perhaps the sixth to the third centuries bce, and presumably were used there – as in Scythia – to mark the burials of leading Scythians deposited in the area. Is this cultural osmosis? We should probably expect osmosis to occur in tandem with the movement of artefacts, so that only good contexts can really answer such questions from case to case. However, the broad pattern is indicated by a range of factors. Particularly notable in this regard is the observable development of a Thraco-Scythian form of what is more familiar as “Scythian animal style,” a term which – it must be understood – already embraces a range of types as we examine the different examples of the style across the great expanse from Siberia to the western Ukraine. As Melyukova observes, Thrace shows both items made in this style among Scythians and, more numerous and more interesting, a Thracian tendency to adapt that style to local tastes, with observable regional distinctions within Thrace itself. Among the Getae and Odrysians the adaptation seems to have been at its height from the later fifth century to the mid-third century (Melyukova 1965, 38; 1979).

The absence of local animal style in Bulgaria before the fifth century bce confirms that we have cultural influences and osmosis at work here, though that is not to say that Scythian tradition somehow dominated its Thracian counterpart, as has been claimed (pace Melyukova 1965, 39; contrast Kitov 1980 and 1984). Of particular interest here is the horse-gear (forehead-covers, cheek-pieces, bridle fittings, and so on) which is found extensively in Romania and Bulgaria as well as in Scythia, both in hoarded deposits and in burials. This exemplifies the development of a regional animal style, not least in silver and bronze, which problematizes the whole issue of the place(s) of its production. Accordingly, the regular designation as “Thracian” of horse-gear from the rich fourth century Scythian burial of Oguz in the Ukraine becomes at least awkward and questionable (further, Fialko 1995). And let us be clear that this is no minor matter, nor even part of a broader debate about the shared development of toreutics among Thracians and Scythians (e.g., Kitov 1980 and 1984). A finely equipped horse of fine quality was a strong statement and striking display of wealth and the power it implied

(…) while Thracian pottery appears at Olbia, Scythian pottery among Thracians is largely confined to the eastern limits of what should probably be regarded as Getic territory, namely the area close to the west of the Dniester, from the sixth century bce. Rather exceptional then is the Scythian pottery noted at Istros, which has been explained as a consequence of the Scythian pursuit of the withdrawing army of Darius and, possibly, a continued Scythian grip on the southern Danube in its aftermath (Melyukova 1965, 34). The archaeology seems to show us, therefore, that the elite Thracians and Scythians were more open to adaptation and acculturation than were their lesser brethren.

palaeo-balkan-languages
Paleo-Balkan languages in Eastern Europe between 5th and 1st century BC. From Wikipedia.

Conclusion

(…) we see distinct peoples and organizations, for example as Sitalces’ forces line up against the Scythians. Much more striking, however, against that general background, are the various ways in which the two peoples and their elites are seen to interact, connect, and share a cultural interface. We see also in Scyles’ story how the Greek cities on the coast of Thrace and Scythia played a significant role in the workings of relationships between the two peoples. It is not simply that these cities straddled the Danube, but also that they could collaborate – witness the honors for Autocles, ca. 300 bce (SEG 49.1051; Ochotnikov 2006) – and were implicated with the interactions of the much greater non-Greek powers around them. At the same time, we have seen the limited reality of familiar distinctions between settled Thracians and nomadic Scythians and the limited role of the Danube too in dividing Thrace and Scythia. The interactions of the two were not simply matters of dynastic politics and the occasional shared taste for artefacts like horse-gear, but were more profoundly rooted in the economic matrix across the region, so that “Scythian” nomadism might flourish in the Dobrudja and “Thracian-style” agriculture and settlement can be traced from Thrace across the Danube as far as Olbia. All of that offers scant justification for the Greek tendency to run together Thracians and Scythians as much the same phenomenon, not least as irrational, ferocious, and rather vulgar barbarians (e.g., Plato, Rep. 435b), because such notions were the result of ignorance and chauvinism. However, Herodotus did not share those faults to any degree, so that we may take his ready movement from Scythians to Thracians to be an indication of the importance of interaction between the two peoples whom he had encountered not only as slaves in the Aegean world, but as powerful forces in their own lands (e.g., Hdt. 4.74, where Thracian usage is suddenly brought into his account of Scythian hemp). Similarly, Thucydides, who quite without need breaks off his disquisition on the Odrysians to remark upon political disunity among the Scythians (Thuc. 2.97, a favorite theme: cf. Hdt. 4.81; Xen., Cyr. 1.1.4). As we have seen throughout this discussion, there were many reasons why Thracians might turn the thoughts of serious writers to Scythians and vice versa.

It seems, following Sikora et al. (2014), that Thracian ‘common’ populations would have more Anatolian Neolithic ancestry compared to more ‘steppe-like’ samples. But there were important differences even between the two nearby samples published from Bulgaria, which may account for the close interaction between Scythians and Thracians we see in Krzewińska et al. (2018), potentially reflected in the differences between the Central, Southern and the South-Central clusters (possibly related to different periods rather than peoples??).

If these R1b-Z2103 were descended from Thracian elites, this would be the first proof of Palaeo-Balkan populations showing mainly R1b-Z2103, as I expect. Their appearance together with haplogroup I2a2a1b1 (also found in Ukraine Neolithic and in the Yamna outlier from Bulgaria) seem to support this regional continuity, and thus a long-lasting cultural and ethnic border roughly around the Danube, similar to the one found in the northern Caucasus.

However, since these samples are some 2,500 years younger than the Yamna expansion to the south, and they are archaeologically Scythians, it is impossible to say. In any case, it would seem that the main expansion of R1a-Z645 lineages to the south of the Danube – and therefore those found among modern Greeks – was mediated by the Slavic expansions centuries later.

krzewinska-scythians-pca
Modified image from Krzewińska et al. (2018), with added Y-DNA haplogroups to each defined Scythian cluster and Sarmatians. Principal component analysis (PCA) plot visualizing 35 Bronze Age and Iron Age individuals presented in this study and in published ancient individuals in relation to modern reference panel from the Human Origins data set. See image with population references.

On the Northern cluster there is a sample of haplogroup R1b-P312 which, given its position on the PCA (apparently even more ‘modern Celtic’-like than the Hallstatt_Bylany sample from Damgaard et al. 2018), it seems that it could be the product of the previous eastward Hallstatt expansion…although potentially also from a recent one?:

Especially important in the archaeology of this interior is the large settlement at Nemirov in the wooded steppe of the western Ukraine, where there has been considerable excavation. This settlement’s origins evidently owe nothing significant to Greek influence, though the early east Greek pottery there (from ca. 650 bce onward: Vakhtina 2007) and what seems to be a Greek graffito hint at its connections with the Greeks of the coast, especially at Olbia, which lay at the estuary of the River Bug on whose middle course the site was located (Braund 2008). The main interest of the site for the present discussion, however, is its demonstrable participation in the broader Hallstatt culture to its west and south (especially Smirnova 2001). Once we consider Nemirov and the forest steppe in connection with Olbia and the other locations across the forest steppe and coastal zone, together with the less obvious movements across the steppe itself, we have a large picture of multiple connectivities in which Thrace bulks large.

scythian-peoples-balkans
Early Iron Age cultures of the Carpathian basin ca. 7-6th century BC, including steppe-related groups. Ďurkovič et al. (2018).

While the above description of clear-cut R1a-Steppe and R1b-Balkans is attractive (and probably more reliable than admixture found in scattered samples of unclear dates), the true ancient genetic picture is more complicated than that:

  • There is nothing in the material culture of the published western Scythians to distinguish the supposed Thracian elites.
  • We have the sample I0575, an Early Sarmatian from the southern Urals (one of the few available) of haplogroup R1b-Z2106, which supports the presence of R1b-Z2103 lineages among Eastern Iranian-speaking peoples.
  • We also have DA30, a Sarmatian of I2b lineage from the central steppes in Kazakhstan (ca. 47 BC – 24 AD).
  • Other Sarmatian samples of haplogroup R remain undefined.
  • There is R1a-Z93 in a late Sarmatian-Hun sample, which complicates the picture of late pastoralist nomads further.

Therefore, the possibility of hidden pockets of Iranian peoples of R1b-Z2103 (maybe also R1b-P312) lineages remains the best explanation, and should not be discarded simply because of the prevalent haplogroups among modern populations, or because of the different clusters found, or else we risk an obvious circular reasoning: “this sample is not (autosomically or in prevalent haplogroups) like those we already had from the steppe, ergo it is not from this or that steppe culture.” Hopefully, the upcoming paper by Järve et al. will help develop a clearer genetic transect of Iranian populations from the steppes.

All in all, the diversity among western Scythians represents probably one of the earliest difficult cases of acculturation to be studied with ancient DNA (obviously not the only one), since Scythians combine unclear archaeological data with limited and conflicting proto-historical accounts (also difficult to contrast with the wide confidence intervals of radiocarbon dates) with different evolving clusters and haplogroups – especially in border regions with strong and continued interactions of cultures and peoples.

With emerging complex cases like these during the Iron Age, I am happy to see that at least earlier expansions show clearer Y-DNA bottlenecks, or else genetics would only add more data to argue about potential cultural diffusion events, instead of solving questions about proto-language expansions once and for all…

Related

Y-chromosome mixture in the modern Corsican population shows different migration layers

mesolithic-europe

Open access Prehistoric migrations through the Mediterranean basin shaped Corsican Y-chromosome diversity, by Di Cristofaro et al. PLOS One (2018).

Interesting excerpts:

This study included 321 samples from men throughout Corsica; samples from Provence and Tuscany were added to the cohort. All samples were typed for 92 Y-SNPs, and Y-STRs were also analyzed.

Haplogroup R represented approximately half of the lineages in both Corsican and Tuscan samples (respectively 51.8% and 45.3%) whereas it reached 90% in Provence. Sub-clade R1b1a1a2a1a2b-U152 predominated in North Corsica whereas R1b1a1a2a1a1-U106 was present in South Corsica. Both SNPs display clinal distributions of frequency variation in Europe, the U152 branch being most frequent in Switzerland, Italy, France and Western Poland. Calibrated branch lengths from whole Y chromosome sequencing [44,45] and ancient DNA studies [46] both indicated that R1a and R1b diversification began relatively recently, about 5 Kya, consistent with Bronze Age and Copper Age demographic expansion. TMRCA estimations are concordant with such expansion in Corsica.

corsica-haplogroups
Spatial frequency maps for haplogroups with frequencies above 3%, their Y-STR based phylogenetic networks in Corsican populations (Blue: North, Green: West, Orange: South, Black: Center and Purple: East) and their TMRCA (in years, +/- SE).

Haplogroup G reached 21.7% in Corsica and 13.3% in Tuscany. Sub-clade G2a2a1a2-L91 accounted for 11.3% of all haplogroups in Corsica yet was not present in Provence or in Tuscany. Thirty-four out of the 37 G2a2a1a2-L91 displayed a unique Y-STR profile, illustrated by the star-like profile of STR networks (Fig 1). G2a2a1a2-L91 and G2a2a-PF3147(xL91xM286) show their highest frequency in present day Sardinia and southern Corsica compared to low levels from Caucasus to Southern Europe, encompassing the Near and Middle East [21,47–50]. Ancient DNA results from Early and Middle Neolithic samples reported the presence of haplogroup G2a-P15 [51–53], consistent with gene flow from the Mediterranean region during the Neolithic transition. Td expansion time estimated by STR for P15-affiliated chromosomes was estimated to be 15,082+/-2217 years ago [49]. Ötzi, the 5,300-year-old Alpine mummy, was derived for the L91 SNP [21]. A genetic relationship between G haplogroups from Corsica and Sardinia is further supported by DYS19 duplication, reported in North Sardinia [14], and observed in the southern part of the Corsica in 9 out of 37 G2a2a1a2-L91 chromosomes and in 4 out of 5 G2a2a-PF3147(xL91xM286) chromosomes, 3 of which displayed an identical STR profile (S4 Table).

This lineage has a reported coalescent age estimated by whole sequencing in Sardinian samples of about 9,000 years ago. This could reflect common ancestors coming from the Caucasus and moving westward during the Neolithic period [48], whereas their continental counterparts would have been replaced by rapidly expanding populations associated with the Bronze Age [46,54,55]. Estimated TMRCA for L91 lineage in Corsica is 4529 +/- 853 years. G-L497 showed high frequencies in Corsica compared to Provence and Tuscany, and this haplogroup was common in Europe, but rare in Greece, Anatolia and the Middle East. Fifteen out of the 17 Corsican G2a2b2a1a1b-L497 displayed a unique Y-STR profile (S4 Table) with an estimated TMRCA of 6867 +/- 1294 years. Haplogroup G2a2b1-M406, associated with Impressed Ware Neolithic markers, along with J2a1-DYS445 = 6 and J2a1b1-M92 [22,49], had very low levels in Corsica. Conversely, G2a2b2a-P303was highly represented and seemed to be independent of the G2a2b1-M406 marker. The 7 G2a2b2a-P303(xL497xM527) Corsican chromosomes displayed a unique Y-STR profile (S4 Table).

pca-corsica
First and second axes of the PCA based on 12 Y-chromosome haplogroup frequencies in 83 west Mediterranean populations.

Haplogroup J, mainly represented by J2a1b-M67(xM92), displayed intermediate frequencies in Corsica compared to Tuscany and Provence. J2a1b-M67(xM92) derived STR network analysis displayed a quite homogeneous profile across the island with an estimated TMRCA of 2381 +/- 449 years (Fig 1) and individuals displaying M67 were peripheral compared to Northwestern Italians (S2 Fig). The haplogroup J2a1-Page55(xM67xM530), characteristic of non-Greek Anatolia [22], was found in the north-west of Corsica. Haplogroup J2a1-DYS445 = 6 was found in the north-west with DYS391 = 10 repeats, and in the far south with DYS391 = 9 repeats, the former was associated with Anatolian Greek samples, whereas the second was found in central Anatolia [22]. The 7 J2b2a-M241 displayed a unique Y-STR profile (S4 Table), they were only detected in the Cap Corse region, this sub-haplogroup shows frequency peaks in both the southern Balkans and northern-central Italy [56] and is associated with expansion from the Near East to the Balkans during Neolithic period [57].

Haplogroup E, mainly represented by E1b1b1a1b1a-V13, displayed intermediate frequencies in Corsica compared to Tuscany and Provence. E1b1b1a1b1a-V13 was thought to have initiated a pan-Mediterranean expansion 7,000 years ago starting from the Balkans [52] and its dispersal to the northern shore of the Mediterranean basin is consistent with the Greek Anatolian expansion to the western Mediterranean [22], characteristic of the region surrounding Alaria, and consistent with the TMRCA estimated in Corsica for this haplogroup. A few E1b1a-V38 chromosomes are also observed in the same regions as V13.

Related:

Haplogroup J spread in the Mediterranean due to Phoenician and Greek colonizations

iron_age_europe_mediterranean

Open access A finely resolved phylogeny of Y chromosome Hg J illuminates the processes of Phoenician and Greek colonizations in the Mediterranean, by Finocchio et al. Scientific Reports (2018) Nº 7465.

Abstract (emphasis mine):

In order to improve the phylogeography of the male-specific genetic traces of Greek and Phoenician colonizations on the Northern coasts of the Mediterranean, we performed a geographically structured sampling of seven subclades of haplogroup J in Turkey, Greece and Italy. We resequenced 4.4 Mb of Y-chromosome in 58 subjects, obtaining 1079 high quality variants. We did not find a preferential coalescence of Turkish samples to ancestral nodes, contradicting the simplistic idea of a dispersal and radiation of Hg J as a whole from the Middle East. Upon calibration with an ancient Hg J chromosome, we confirmed that signs of Holocenic Hg J radiations are subtle and date mainly to the Bronze Age. We pinpointed seven variants which could potentially unveil star clusters of sequences, indicative of local expansions. By directly genotyping these variants in Hg J carriers and complementing with published resequenced chromosomes (893 subjects), we provide strong temporal and distributional evidence for markers of the Greek settlement of Magna Graecia (J2a-L397) and Phoenician migrations (rs760148062). Our work generated a minimal but robust list of evolutionarily stable markers to elucidate the demographic dynamics and spatial domains of male-mediated movements across and around the Mediterranean, in the last 6,000 years.

greek-phoenician
J2-L397. The star indicates the centroid of derived alleles. The solid square indicates the centroid of ancestral alleles, with its 95% C.I. (ellipse). In the insets: distributions of the pairwise sampling distances (in Km) for the carriers of the ancestral (black) and derived (white) allele, with solid and dashed lines indicating the respective averages. At right: median joining network of 7-STR haplotypes and SNPs in the same groups, with sectors coloured according to sampling location. Haplotype structure is detailed for some nodes, in the order YCA2a-YCA2b-DYS19-DYS390-DYS391-DYS392-DYS393 (in italics).

Interesting excerpts:

Two features of our tree are at odds with the simplistic idea of a dispersal of Hg J as a whole from the Middle East towards Greece and Italy and an accompanying radiation26. First, there is little evidence of sudden diversification between 15 and 5 kya, a period of likely population increase and pressure for range expansion, due to the Agricultural revolution in the Fertile Crescent. Second, within each subclade, lineages currently sampled in Turkey do not show up as preferentially ancestral. Both findings are replicated and reinforced by examining the previous landmark studies. Our Turkish samples do not coalesce preferentially to ancestral nodes when mapped onto these studies’ trees.

Additional relevant information on the entire Hg J comes from the discontinuous distribution of J2b-M12. The northern fringe of our sample is enriched in the J2b-M241 subclade, which reappears in the gulf of Bengal38,45, with low frequencies in the intervening Iraq46 and Iran47. No J2b-M12 carriers were found among 35 modern Lebanese, as contrasted to one of two ancient specimens from the same region35.

In summary, a first conclusion of our sequencing effort and merge with available data is that the phylogeography of Hg J is complex and hardly explained by the presence of a single population harbouring the major lineages at the onset of agriculture and spreading westward. A unifying explanation for all the above inconsistencies could be a centre of initial radiation outside the area here sampled more densely, i.e. the Caucasus and regions North of it, from which different Hg J subclades may have later reached mainland Italy, Greece and Turkey, possibly following different routes and times. Evidence in this direction comes from the distribution of J2a-M41045,48 and the early-49 or mid-Holocene50 southward spread of J1.

greek-colonization
Supplemental Figure 7. Maps of sampling locations for the carriers of the derived allele (white triangle point down) at the indicated SNP vs carriers of the ancestral allele (black triangle point-up), conditioned on identical genotype at the same most terminal marker. Coastlines were drawn with the R packages18 “map” and “mapproj” v. 3.1.3 (https://cran.r-project.org/web/packages/mapproj/index.html), and additional features added with default functions. The star triangle indicates the centroid of derived alleles. The solid square indicates the centroid of ancestral alleles, with its 95% C.I. (ellipse). In the insets: distributions of the pairwise sampling distances (in Km) for the carriers of the ancestral (black) and derived (white) allele, with solid and dashed lines indicating the respective averages. At right: median joining network of 7-STR haplotypes and SNPs in the same groups, with sectors coloured according to sampling location. Haplotype structure is detailed for some nodes, in the order YCA2a-YCA2b-DYS19-DYS390-DYS391-DYS392-DYS393 (in italics).

The lineage defined by rs779180992, belonging to J2b-M205, and dated at 4–4.5 kya, has a radically different distribution, with derived alleles in Continental Italy, Greece and Northern Turkey, and two instances in a Palestinian and a Jew. The interpretation of the spread of this lineage is not straightforward. Tentative hypotheses are linked to Southward movements that occurred in the Balkan Peninsula from the Bronze Age29,53, through the Roman occupation and later54.

The slightly older (5.6–6.3 kya) branch 98 lineage displays a similar trend of a Eastward positioning of derived alleles, with the notable difference of being present in Sardinia, Crete, Cyprus and Northern Egypt. This feature and the low frequency of the parental J2a-M92 lineage in the Balkans27 calls for an explanation different from the above.

Finally, we explored the distribution of J2a-L397 and three derived lineages within it. J2a-L397 is tightly associated with a typical DYS445 6-repeat allele. This has been hypothesized as a marker of the Greek colonizations in the Mediterranean55, based on its presence in Greek Anatolia and Provence (France), a region with attested Iron Age Greek contribution. All of our chromosomes in this clade were characterized also by DYS391(9), confirming their Anatolian Greek signature. We resolved the J2a-L397 clade to an unprecedented precision, with three internal markers which allow a finer discrimination than STRs. The ages of the three lineages (2.0–3.0 kya) are compatible with the beginning of the Greek colonial period, in the 8th century BCE. The three subclades have different distributions (Fig. 2B), with two (branches 57, 59) found both East and West to Greece, and one only in Italy (branch 58). As to Mediterranean Islands, J2a-L397 was found in Cyprus56 and Crete43. Its presence as one of the three branches 57–59 will represent an important test. In Italy all three variants were found mainly along the Western coast (18/25), which hosted the preferred Greek trade cities. The finding of all three differentiated lineages in Locri excludes a local founder effect of a single genealogy. Interestingly, an important Greek colony was established in this location, with continuity of human settlement until modern times. The sample composed of the same subjects displayed genetic affinities with Eastern Greece and the Aegean also at autosomal markers57. In summary, the distributions of branches 57–59 mirror the variety of the cities of origin and geographic ranges during the phases of the colonization process58.

So, there you have it, another proof that haplogroup J and CHG-related ancestry in the Mediterranean was mainly driven by different (and late) expansions of historic peoples.

Related:

Pre-Roman and Roman mitogenomes from Southern Italy

vagnari-cemetery-haplogroups-superimposed

Ph.D. thesis Assessing Migration and Demographic Change in pre-Roman and Roman Period Southern Italy Using Whole-Mitochondrial DNA and Stable Isotope Analysis, or The Biogeographic Origins of Iron Age Peucetians and Working-Class Romans From Southern Italy, by Matthew Emery, McMaster University (2018).

Abstract (emphasis mine):

Assessing population diversity in southern Italy has traditionally relied on archaeological and historic evidence. Although informative, these lines of evidence do not establish specific instances of within lifetime mobility, nor track population diversity over time. In order to investigate the population structure of ancient South Italy I sequenced the mitochondrial DNA (mtDNA) from 15 Iron Age (7th – 4th c. BCE) and 30 Roman period (1st – 4th c. BCE) individuals buried at Iron Age Botromagno and Roman period Vagnari, in southern Italy, and analyzed δ18O and 87Sr/86Sr values from a subset of the Vagnari skeletal assemblage.

Phylogenetic analysis of 15 Iron Age mtDNAs together with 231 mtDNAs spanning European prehistory suggest that southern Italian Iapygians share close genetic affinities to Neolithic populations from eastern Europe and the Near East. Population pairwise analysis of Iron Age, Roman, and mtDNA datasets spanning the pan-Mediterranean region (n=357), indicate that Roman maternal genetic diversity is more similar to Neolithic and Bronze Age populations from central Europe and the eastern Mediterranean, respectively, than to Iron Age Italians. Genetic distance between population age categories imply moderate mtDNA turnover and constant population size during the Roman conquest of South Italy in the 3rd century BCE.

In order to determine the local versus non-local demographic at Vagnari, I measured the 87Sr/86Sr and 18O/16O of composition of 43 molars, and the 87Sr/86Sr composition of an additional 13 molars, and constructed a preliminary 87Sr/86Sr variation map of the Italian peninsula using disparate 87Sr/86Sr datasets. The relationship between 87Sr/86Sr and previously published δ18O data suggest a relatively low proportion of migrants lived at Vagnari (7%).

This research is the first to generate whole-mitochondrial DNA sequences from Iron Age and Roman period necropoleis, and demonstrates the ability to gain valuable information from the integration of aDNA, stable isotope, archaeological and historic evidence.

italy-iron-age-mtdna
mtDNA haplogroup composition between Botromagno (7th – 4th century BCE; n=15) and Vagnari (1st – 4th century CE; n=30) skeletal assemblages.

Interesting excerpts:

Taken together, population pairwise ΦST, and the distribution of mtDNA haplotypes in relation to the comparative mtDNA data set show that the Iron Age southern Italians likely descended from early to late Neolithic farmers from Anatolia and possibly as far East as the Caucasus, and from migrants arriving from eastern Europe around the late Neolithic/early Bronze Age. These findings support previous hypotheses that the ancestors of the Iapygians may have originated in the eastern Balkan region, or derive shared ancestry with a common source population from eastern Europe. Alternatively, southern Italian Iron Age mtDNA variation might also reflect LGM gene flow between southwestern European, Mediterranean, and Carpathian basin refugia, which was suggested for haplogroup subclusters of U5 and J (Malyarchuk et al., 2010; Pala et al., 2012). Future mtDNA (and nuclear DNA) analysis comprised of a larger Iron Age data set from southern Italy is necessary to answer Theodor Mommsen’s initial hypothesis that the Iapygians were the oldest immigrants to the southern Italian region.

Our investigation provides the first mtDNA evidence for the maternal ancestral affiliations of a subset of the Iapygian individuals recovered from southern Italy, and suggests a closer genetic link to European Neolithic and Iron Age Armenians, than to Bronze Age Aegeans. Future comparative ancient DNA data using whole-genome SNP, mtDNA, and NRY-chromosome analysis of pre-Roman populations will provide complementary evidence for the ancestral roots of understudied Iron Age individuals from Italy.

Illyrian_colonies_in_Italy_550_BC
Simplistic map of Illyrian colonies in Italy 550 BCE, from Wikipedia

Archaeological evidence indicates that the Iapygians traded and incorporated Hellenistic elements into their material and cultural traditions (Small, 1992; Peruzzi, 2016). These changes are most apparent in burial custom and ceramic production, and become increasingly prominent by 2400 BP (Peruzzi, 2016). Further evidence shows that Iron Age communities across South Italy retracted in size amidst ongoing conflict between colonies in Magna Graecia, and Rome and Carthage (Small, 1992). This apparent change was interpreted as a decline in local populations throughout the region. However, Bayesian Skygrid analysis using the mtDNA profiles of 15 Iapygians and 30 Roman period individuals suggest that female effective population size was comparable between the two populations. In Chapter 4, population distance (measured as population pairwise ΦST values) across a range of mtDNAs obtained from the pan-Mediterranean, European, and western Asian regions suggest closer maternal affinities to Neolithic and Bronze Age populations from the eastern Mediterranean as a cohort, than with Iron Age Italians. This finding points to moderate mtDNA turnover, and is likely the consequence of Roman gene flow stemming from central and northern Italy via the migration and subsequent occupation by Roman colonies after 2250 BP.

Roman Imperial pursuits peaked by ~2050 BP. This extension of power, coupled with an increase in food and materials procurement, was driven by a substantial labour force comprised of both low status Romans and slaves (Harris, 1980; Bradley, 1987, 1994, 2000). Although several attempts have been made to quantify the number of slaves required to maintain the Roman economy, it is unknown what fraction of the Roman population was slave-owned (~approximately 1 to 3 million by 2050 BP) (Scheidel, 2005). Rome’s slave acquisition during the early centuries of the Republic was likely maintained through military campaigns and conquest, a trend that is well documented in Italy (Scheidel, 1997, 1999, 2005; Harris, 1999; Small, 2002). However, once territory was secured, local slave populations were likely maintained through one or a combination of the following: i) the importation of slaves from non-local regions, ii) were born to slave-owned parents, or iii) were voluntarily self-enslaved to acquire subsistence (Harris, 1999). The importation of foreign slaves was likely more costly than maintaining a self-reproducing slave population, especially in rural areas. As such, rural Roman necropoleis, such Vagnari, provide an opportune case to determine the local versus non-local demographic. Archaeological evidence suggests that Vagnari was involved in agriculture and industrial procurement, and was likely staffed by low-class individuals possibly including slaves (Small et al., 2000). However, without direct archaeological or epigraphic evidence, it is impossible to identify the proportion of slaves at rural sites.

italy-iron-age-roman-plot
Multi-dimensional scaling plots showing pairwise ΦST values by a) age and b) country. We removed age and geographic categories with less than 5 mtDNA sequence representation to reduce scaling stress, which decreased the sample size from 402 mtDNAs to n = 378 by age, and n= 382 by country. a) MDS plot of the mtDNA categorized by country of origin; b) MDS of mtDNA dataset by age spanning the Upper Paleolithic (pre-LGM) to the Roman period. IronAge 1 = Italian Iron Age samples; IronAge 2 = Armenian Iron Age samples; Roman 1 = Italian Roman samples; Roman 2 = Egyptian Roman samples; TIP = Third Intermediary Period (Egypt); LP = Late Period (Egypt); PP = Ptolemaic Period (Egypt).

(…) The isotope values presented in Chapter 3 obtained from 56 Roman individuals buried at Vagnari suggest that over half (58%) were born directly at Vagnari, with a further 34% originating from South Italy. Only 7% (3/43 with both δ18O and 87Sr/86Sr values) of the individuals sampled resulted in isotope values non-local to the southern peninsula. Two of these individuals originated from either northern Italy or, more broadly, from central Europe, while one individual likely originated from North Africa. Overall, the isotope data suggest a low number of immigrants at Vagnari, which conforms with the population pairwise (ΦST) data for the Iron Age and Roman mtDNAs, and suggests that as the Romans occupied the region, they populated their Imperial properties with people from central Italy (possible the region of Latium, and the surrounding environs of Rome). These results also integrate with the historical evidence concerning the Roman slave economy during the Imperial period. Future research using a larger comparative dataset comprised of pre-Roman and Roman period mtDNAs, δ18O and, 87Sr/86Sr results will refine the interpretations outlined here.

A paper from this thesis is already published in a peer-review journal, Mapping the origins of Imperial Roman workers (1st–4th century CE) at Vagnari, Southern Italy, using 87Sr/86Sr and δ18O variability, Am J Phys Anthropol (2018).

Related:

Spatio-temporal deixis and cognitive models in early Indo-European

helios-rising

Interesting article, Spatio-temporal deixis and cognitive models in early Indo-European, by Annamaria Bartolotta, Cognitive Linguistics (2018); 29(1):1-44.

Abstract (emphasis mine):

This paper is a comparative study based on the linguistic evidence in Vedic Sanskrit and Homeric Greek, aimed at reconstructing the space-time cognitive models used in the Proto-Indo-European language in a diachronic perspective. While it has been widely recognized that ancient Indo-European languages construed earlier (and past) events as in front of later ones, as predicted in the Time-Reference-Point mapping, it is less clear how in the same languages the passage took place from this ‘archaic’ Time-RP model or non-deictic sequence, in which future events are behind or follow the past ones in a temporal sequence, to the more recent ‘post-archaic’ Ego-RP model that is found only from the classical period onwards, in which the future is located in front and the past in back of a deictic observer. Data from the Rigveda and the Homeric poems show that an Ego-RP mapping with an ego-perspective frame of reference (FoR) could not have existed yet at an early Indo-European stage. In particular, spatial terms of front and behind turn out to be used with reference not only to temporal events, but also to east and west respectively, thus presupposing the existence of an absolute field-based FoR which the temporal sequence is metaphorically related to. Specifically, sequence is relative position on a path appears to be motivated by what has been called day orientation frame, in which the different positions of the sun during the day motivate the mapping of front onto ‘earlier’ and behind onto ‘later’, without involving ego’s ‘now’. These findings suggest that early Indo-European still had not made use of spatio-temporal deixis based on the tense-related ego-perspective FoR found in modern languages.

Featured image, from the article: Helios rising from the sea (blacas red-figured calyx-krater, fifth century B.C., British Museum). Related quote from the article:
“Interestingly, the archeological evidence supports that time could be spatialized along the lateral axis. In ancient Greek art the sun is represented as moving from right to left. Such orientation can be observed, for instance, on the Blacas red-figured calyx-krater of the fifth century B.C. (London, British Museum), where Helios is found at the extreme right of the scene and proceeds to the viewer’s left, following Eos, i.e., the dawn.”

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