A Song of Sheep and Horses, revised edition, now available as printed books

cover-song-sheep-and-horses

As I said 6 months ago, 2019 is a tough year to write a blog, because this was going to be a complex regional election year and therefore a time of political promises, hence tenure offers too. Now the preliminary offers have been made, elections have passed, but the timing has slightly shifted toward 2020. So I may have the time, but not really any benefit of dedicating too much effort to the blog, and a lot of potential benefit of dedicating any time to evaluable scientific work.

On the other hand, I saw some potential benefit for publishing texts with ISBNs, hence the updates to the text and the preparation of these printed copies of the books, just in case. While Spain’s accreditation agency has some hard rules for becoming a tenured professor, especially for medical associates (whose years of professional experience are almost worthless compared to published peer-reviewed papers), it is quite flexible in assessing one’s merits.

However, regional and/or autonomous entities are not, and need an official identifier and preferably printed versions to evaluate publications, such as an ISBN for books. I took thus some time about a month ago to update the texts and supplementary materials, to publish a printed copy of the books with Amazon. The first copies have arrived, and they look good.

series-song-sheep-horses-cover

Corrections and Additions

Titles
I have changed the names and order of the books, as I intended for the first publication – as some of you may have noticed when the linguistic book was referred to as the third volume in some parts. In the first concept I just wanted to emphasize that the linguistic work had priority over the rest. Now the whole series and the linguistic volume don’t share the same name, and I hope this added clarity is for the better, despite the linguistic volume being the third one.

Uralic dialects
I have changed the nomenclature for Uralic dialects, as I said recently. I haven’t really modified anything deeper than that, because – unlike adding new information from population genomics – this would require for me to do a thorough research of the most recent publications of Uralic comparative grammar, and I just can’t begin with that right now.

Anyway, the use of terms like Finno-Ugric or Finno-Samic is as correct now for the reconstructed forms as it was before the change in nomenclature.

west-east-uralic-schema

Mediterranean
The most interesting recent genetic data has come from Iberia and the Mediterranean. Lacking direct data from the Italian Peninsula (and thus from the emergence of the Etruscan and Rhaetian ethnolinguistic community), it is becoming clearer how some quite early waves of Indo-Europeans and non-Indo-Europeans expanded and shrank – at least in West Iberia, West Mediterranean, and France.

Finno-Ugric
Some of the main updates to the text have been made to the sections on Finno-Ugric populations, because some interesting new genetic data (especially Y-DNA) have been published in the past months. This is especially true for Baltic Finns and for Ugric populations.

ananino-culture-new

Balto-Slavic
Consequently, and somehow unsurprisingly, the Balto-Slavic section has been affected by this; e.g. by the identification of Early Slavs likely with central-eastern populations dominated by (at least some subclades of) hg. I2a-L621 and E1b-V13.

Maps
I have updated some cultural borders in the prehistoric maps, and the maps with Y-DNA and mtDNA. I have also added one new version of the Early Bronze age map, to better reflect the most likely location of Indo-European languages in the Early European Bronze Age.

As those in software programming will understand, major changes in the files that are used for maps and graphics come with an increasing risk of additional errors, so I would not be surprised if some major ones would be found (I already spotted three of them). Feel free to communicate these errors in any way you see fit.

bronze-age-early-indo-european
European Early Bronze Age: tentative langage map based on linguistics, archaeology, and genetics.

SNPs
I have selected more conservative SNPs in certain controversial cases.

I have also deleted most SNP-related footnotes and replaced them with the marking of each individual tentative SNP, leaving only those footnotes that give important specific information, because:

  • My way of referencing tentative SNP authors did not make it clear which samples were tentative, if there were more than one.
  • It was probably not necessary to see four names repeated 100 times over.
  • Often I don’t really know if the person I have listed as author of the SNP call is the true author – unless I saw the full SNP data posted directly – or just someone who reposted the results.
  • Sometimes there are more than one author of SNPs for a certain sample, but I might have added just one for all.
ancient-dna-all
More than 6000 ancient DNA samples compiled to date.

For a centralized file to host the names of those responsible for the unofficial/tentative SNPs used in the text – and to correct them if necessary -, readers will be eventually able to use Phylogeographer‘s tool for ancient Y-DNA, for which they use (partly) the same data I compiled, adding Y-Full‘s nomenclature and references. You can see another map tool in ArcGIS.

NOTE. As I say in the text, if the final working map tool does not deliver the names, I will publish another supplementary table to the text, listing all tentative SNPs with their respective author(s).

If you are interested in ancient Y-DNA and you want to help develop comprehensive and precise maps of ancient Y-DNA and mtDNA haplogroups, you can contact Hunter Provyn at Phylogeographer.com. You can also find more about phylogeography projects at Iain McDonald’s website.

Graphics
I have also added more samples to both the “Asian” and the “European” PCAs, and to the ADMIXTURE analyses, too.

I previously used certain samples prepared by amateurs from BAM files (like Botai, Okunevo, or Hittites), and the results were obviously less than satisfactory – hence my criticism of the lack of publication of prepared files by the most famous labs, especially the Copenhagen group.

Fortunately for all of us, most published datasets are free, so we don’t have to reinvent the wheel. I criticized genetic labs for not releasing all data, so now it is time for praise, at least for one of them: thank you to all responsible at the Reich Lab for this great merged dataset, which includes samples from other labs.

NOTE. I would like to make my tiny contribution here, for beginners interested in working with these files, so I will update – whenever I have time – the “How To” sections of this blog for PCAs, PCA3d, and ADMIXTURE.

-iron-age-europe-romans
Detail of the PCA of European Iron Age populations. See full versions.

ADMIXTURE
For unsupervised ADMIXTURE in the maps, a K=5 is selected based on the CV, giving a kind of visual WHG : NWAN : CHG/IN : EHG : ENA, but with Steppe ancestry “in between”. Higher K gave worse CV, which I guess depends on the many ancient and modern samples selected (and on the fact that many samples are repeated from different sources in my files, because I did not have time to filter them all individually).

I found some interesting component shared by Central European populations in K=7 to K=9 (from CEU Bell Beakers to Denmark LN to Hungarian EBA to Iberia BA, in a sort of “CEU BBC ancestry” potentially related to North-West Indo-Europeans), but still, I prefer to go for a theoretically more correct visualization instead of cherry-picking the ‘best-looking’ results.

Since I made fun of the search for “Siberian ancestry” in coloured components in Tambets et al. 2018, I have to be consistent and preferred to avoid doing the same here…

qpAdm
In the first publication (in January) and subsequent minor revisions until March, I trusted analyses and ancestry estimates reported by amateurs in 2018, which I used for the text adding my own interpretations. Most of them have been refuted in papers from 2019, as you probably know if you have followed this blog (see very recent examples here, here, or here), compelling me to delete or change them again, and again, and again. I don’t have experience from previous years, although the current pattern must have been evidently repeated many times over, or else we would be still talking about such previous analyses as being confirmed today…

I wanted to be one step ahead of peer-reviewed publications in the books, but I prefer now to go for something safe in the book series, rather than having one potentially interesting prediction – which may or may not be right – and ten huge mistakes that I would have helped to endlessly redistribute among my readers (online and now in print) based on some cherry-picked pairwise comparisons. This is especially true when predictions of “Steppe“- and/or “Siberian“-related ancestry have been published, which, for some reason, seem to go horribly wrong most of the time.

I am sure whole books can be written about why and how this happened (and how this is going to keep happening), based on psychology and sociology, but the reasons are irrelevant, and that would be a futile effort; like writing books about glottochronology and its intermittent popularity due to misunderstood scientist trends. The most efficient way to deal with this problem is to avoid such information altogether, because – as you can see in the current revised text – they wouldn’t really add anything essential to the content of these books, anyway.

Continue reading

Official site of the book series:
A Song of Sheep and Horses: eurafrasia nostratica, eurasia indouralica

More Hungarian Conquerors of hg. N1c-Z1936, and the expansion of ‘Altaic-Uralic’ N1c

Open access Y-chromosomal connection between Hungarians and geographically distant populations of the Ural Mountain region and West Siberia, by Post et al. Scientific Reports (2019) 9:7786.

Hungarian Conquerors

More interesting than the study of modern populations of the paper is the following excerpt from the introduction, referring to a paper that is likely in preparation, Európai És Ázsiai Apai Genetikai Vonalak A Honfoglaló Magyar Törzsekben, by Fóthi, E., Fehér, T., Fóthi, Á. & Keyser, C., Avicenna Institute of Middle Eastern Studies (2019):

Certain chr-Y lineages from haplogroup (hg) N have been proposed to be associated with the spread of Uralic languages. So far, hg N3 has not been reported for Indo-European speaking populations in Central Europe, but it is present among Hungarians, although the proportion of hg N in the paternal gene pool of present-day Hungarians is only marginal (up to 4%) compared to other Uralic speaking populations. It has been shown earlier that one of the sub-clades of hg N – N3a4-Z1936 – could be a potential link between two Ugric speaking populations: the Hungarians and the Mansi. It is also notable that some ancient Hungarian samples from the 9th and 10th century Carpathian Basin belonged to this hg N sub-clade: Three Z1936 samples were found in the Upper-Tisza area (Karos II, Bodrogszerdahely/Streda nad Bodrogom) and two in the Middle-Tisza basin cemeteries (Nagykörű and Tiszakécske). The haplotype of the Nagykörű sample is identical with one contemporary Hungarian sample from Transylvania that tested positive for B545 marker downstream of N3a4-Z193632. Similar findings come from the maternal gene pool of historical Hungarians: the analyses of early medieval aDNA samples from Karos-Eperjesszög cemeteries revealed the presence of mtDNA hgs of East Asian provenance.

A commenter recently wrote that in a study by Fehér (probably this one) two Hungarian conquerors, from Ormenykut and Tuzser, will be of hg. N1c-2110. Assuming no other lineages will appear, this would leave the proportion of N1c-L392 vs. R1a-Z280/Z93 closer to the reported proportion of hg. N vs. R1a (5 vs. 2) among Sargat samples, and is thus compatible with a direct migration of Hungarians from around the Urals.

However, the sampling of Iron Age populations around the Urals is scarce, and we don’t know what other lineages these studied Magyars will have, but – based on the known variability of the published ones, and on the ca. 50-60 early Magyar males available to date in previous studies to obtain Y-chromosome haplogroups – I would say these reported N1c lineages are just a tiny proportion of what’s to come…

“Altaic-Uralic” N1c

altaic-uralic-n1c-haplogroup
Phylogenetic tree of hg N3a4. Phylogenetic tree of 33 high coverage Y-chromosomes from
haplogroup N3a4 was reconstructed with BEAST v.1.7.5 software package.

Archaeogenetic studies based on mtDNA haplotypes have shown that ancient Hungarians were relatively close to contemporary Bashkirs who are a Turkic speaking population residing in the Volga-Ural region. Another study reported excessive identical-by-descent (IBD) genomic segments shared between the Ob-Ugric speaking Khantys and Bashkirs but a moderate IBD sharing between Turkic speaking Tatars and their neighbours including Bashkirs.

Phylogenetic tree of hg N3a4 has two main sub-clades defined by markers B535 and B539 that diverged around 4.9 kya (95% confidence interval [CI] = 3.7–6.3 kya). Inner sub-clades of N3a4-B539 (defined by markers B540 and B545) split 4.2 kya (95% CI = 3.0–5.6 kya). (…) The phylogenetic tree reveals that all five Hungarian samples belong to N3a4-B539 sub-clade that they share with Ob-Ugric speaking Khanty and Mansi, and Turkic speaking Bashkirs and Tatars from the Volga-Ural region. Hungarian and Bashkir chrY lineages belong to both sub-clades of N3a4-B539.

Modern distribution of the “Ugric N1c”

To test the presence and proportions of hg N3a4 lineages in a more comprehensive sample set and with a higher phylogenetic resolution level compared to earlier studies, we analysed the genotyping data of about 5000 Eurasian individuals, including West Siberian Mansi and Khanty who are linguistically closest to Hungarians

n3a4-n1c-z1936-ugric
Map of the entire hg N3a4.

There is a clear difference in geographic distribution patterns of these two hg N3a4 sub-clades. Hg N3a4-B535 (Fig. 3b) is common mostly among Finnic (Finns, Karelians, Vepsas, Estonians) and Saami speaking populations in North eastern Europe. The highest frequency is detected in Finns (~44%) but it also reaches up to 32% in Vepsas and around 20% in Karelians, Saamis and North Russians. The latter are known to have changed their language or to be an admixed population with reported similar genetic composition to their Finnic speaking neighbors. The frequency of N3a4-B535 rapidly decreases towards south to around 5% in Estonians, being almost absent in Latvians (1%) and not found among Lithuanians. Towards east its frequency is from 1–9% among Eastern European Russians and populations of the Volga-Ural region such as Komis, Mordvins and Chuvashes (…)

n3a4-n1c-z1936-finnic-samic
Map of N3a4 subclades defined by B535.

Hg N3a4-B539, on the other hand, is prevalent among Turkic speaking Bashkirs and also found in Tatars but is entirely missing from other populations of the Volga-Ural region such as Uralic speaking Udmurts, Maris, Komis and Mordvins, and in Northeast Europe, where instead N3a4-B535 lineages are frequent. Besides Bashkirs and Tatars in Volga-Ural region, N3a4-B539 is substantially represented in West Siberia among Ugric speaking Mansis and Khantys. Among Hungarians, however, N3a4-B539 has a subtle frequency of 1–4%.

n3a4-n1c-z1936-ugric-bashkir
Map of N3a4 subclades defined by B539, with a local snapshot showing the N3a4-B539 distribution among Hungarian speakers.

The battle to appropriate N1c-L392

So, basically, the team of Kristiina Tambets is arguing that N1c-VL29 expanded Finnic to the East Baltic (hence from a common Finno-Mordvinic dialect splitting ca. 600 BC on?) because, you know, apparently the agreed separation of known Uralic dialects from ca. 2000 BC, and their Bronze Age presence around the Baltic, is not valid when you follow haplogroups instead of languages or archaeology.

But now this other group of Tambets (co-author of this paper) considers that hg. N1c-Z1936 – which is probably behind the N1c-L392 samples from Lovozero Ware in the Kola Peninsula – represent either the True Uralic-speaking Palaeo-Arctic peoples, or else merely Ugric-speaking peoples which happened to expand to Fennoscandia but left no trace of their language…

To accept this identification you only have to NOT ask why:

  • N1c is first found in ancient cultures close to Lake Baikal.
  • N1c-L392 appears in ancient East Asian populations speaking completely different languages, with Altaic and Uralic being just some among many Palaeo-Siberian populations where the haplogroup will pop up.
  • Turkic populations like Bashkirs and Tatars (who expanded to the Volga through the southern Urals before the expansion of Hungarians) show a shared distribution of the B539 haplotype with Hungarians.
  • The phylogenetic tree and areas of N1c-L392 expansions don’t make any sense in light of the known linguistic and cultural expansions of Uralic-speaking peoples.

In fact, the Hungarian research group of Neparáczki – publishing the recent paper on Hungarian Conquerors – was apparently looking for a connection with Turkic peoples to support some traditional Turanian myths, and they found it in some scattered R1a-Z93 samples which supposedly connect Hungarian Conquerors to Huns (?), instead of looking for this closer link through N1c-Z1936 (especially haplotype B539)…

Also, is it me or are there two opposed trends with completely different interpretations among researchers publishing papers about hg. N1c: one systematically arguing for Altaic origins, and another for Uralic ones?

If somebody sees some complex reasoning behind the discussions of all these recent papers, beyond the simplest “let’s follow N for Uralic/Altaic”, feel free to comment below. Just so I can understand what I might be doing wrong in assessing Neolithic and Bronze Age migrations in linguistics and archaeology with help of ancient haplogroups coupled with ancestral components, but these researchers are doing right by playing with obsessive ideas born out of the 2000s coupled with phylogenetic trees and maps of modern haplogroup distributions…

This is probably going to be this blog’s most used image in 2019:

horse-meme-steppe-ancestry

Related

“Dinaric I2a” and the expansion of Common Slavs from East-Central Europe

late-iron-age-eastern-europe

A recently published abstract for an upcoming chapter about Early Slavs shows the generalized view among modern researchers that Common Slavs did not spread explosively from the east, an idea proper of 19th-century Romantic views about ancestral tribes of pure peoples showing continuity since time immemorial.

Migrations and language shifts as components of the Slavic spread, by Lindstedt and Salmela, In: Language contact and the early Slavs, Eds. Tomáš Klír, Vít Boček, Universitätsverlag Winter (2019):

The rapid spread of the Proto-Slavic language in the second half of the first millennium CE was long explained by the migration of its speakers out of their small primary habitat in all directions. Starting from the 1980s, alternative theories have been proposed that present language shift as the main scenario of the Slavic spread, emphasizing the presumed role of Slavic as the lingua franca of the Avar Khaganate. Both the migration and the language shift scenarios in their extreme forms suffer from factual and chronological inaccuracy. On the basis of some key facts about human population genetics (the relatively recent common ancestry of the East European populations), palaeoclimatology (the Late Antique Little Ice Age from 536 to around 660 CE), and historical epidemiology (the Justinianic Plague), we propose a scenario that includes a primary rapid demographic spread of the Slavs followed by population mixing and language shifts to and from Slavic in different regions of Europe. There was no single reason for the Slavic spread that would apply to all of the area that became Slavic-speaking. The northern West Slavic area, the East Slavic area, and the Avar sphere and South-Eastern Europe exhibit different kinds of spread: mainly migration to a sparsely populated area in the northwest, migration and language shift in the east, and a more complicated scenario in the southeast. The remarkable homogeneity of Slavic up to the jer shift was not attributable to a lingua-franca function in a great area, as is often surmised. It was a founder effect: Proto-Slavic was originally a small Baltic dialect with little internal variation, and it took time for the individual Slavic languages to develop in different directions.

While I would need to read the whole chapter, in principle it seems easier to agree with this summary than with Curta’s (sort of diffuse) Danubian origin of Common Slavic, based on the likely origin of the Balto-Slavic expansion with the Trzciniec and/or Lusatian culture, close to the Baltic.

A multi-ethnic Chernyakhov culture

In a sneak peek to the expected Järve et al. (2019) paper in review, there are three Chernyakhov samples (ca. calAD 350-550) with different ancestry probably corresponding to the different regions where they stem from (see image below), which supports the idea that Iron Age eastern Europe was a true melting pot where the eventual language of the different cultures depended on many different factors:

chernyakhov-samples-region
Map of the samples from Järve et al. (2019).

From the paper:

The Chernyakhiv culture was likely an ethnically heterogeneous mix based on Goths (Germanic tribes) but also including Sarmatians, Alans, Slavs, late Scythians and Dacians – the entire ancient population of the northern coast of the Black Sea.

Contacts with neighbouring regions were active, and the Chernyakhiv culture is associated with a number of historical events that took place in Europe at that time. In particular, during the Scythian or Gothic wars of the 230s and 270s, barbarians living in the territory of the Chernyakhiv culture (Goths, Ferules, Carps, Bastarns, etc.) carried out regular raids across the Danube Limes of the Roman Empire. However, from the end of the 3rd century the relations of the barbarians with the Roman Empire gained a certain stability. From the reign of Constantine I the Goths, who were part of the Chernyakhiv culture, became federates (military allies) of the Empire.

The Goths also interacted with the inhabitants of the East European forest zone. The Roman historian Jordanes described the military campaigns of the Gothic king Ermanaric against northern peoples (the ancestors of Vends, Slavs, etc., and the inhabitants of the northern Volga region).

NOTE. As it has become traditional in writings about eastern Europe, ‘Slavs’ are assumed – for no particular reason – to be part of the ‘northern peoples of the forest’ since who knows when exactly, and thus appear mentioned in this very text simultaneously as part of Chernyakhov, but also part of peoples to the north of Chernyakhov warring against them…

admixture-chernyakhov
Proportions of Eastern Hunter-Gatherer (EHG, blue), Natufian (red) and Altaian (green) ancestries in Scythian/Sarmatian groups and groups pre- and postdating them inferred using the a) qpAdm and b) ChromoPainter/NNLS method. c–e Correlation of qpAdm and CP/NNLS proportions for the three putative sources evaluated. Steppe populations predating the Scythians: Yamnaya_Ukraine [26], Yamnaya_Kalmykia [15], Ukr_BA (this study). Scythians and Sarmatians: Nomad_IA [15], Scythian_East and Sarmatian_SU [3], Hungarian Scythian, Sarmatian, Central Saka, Tian Shan Saka and Tagar [1], Scy_Ukr, ScySar_SU and Scy_Kaz (this study). Population postdating the Scythians: Chern (this study). See also Table S3.

Genetic variation

(…) the Chernyakhiv samples overlapped with modern Europeans, representing the most ‘western’ range of variation among the groups of this study.

After the end of the Scythian period in the western Eurasian Steppe, the Chernyakhiv culture samples have higher Near Eastern affinity compared to the Scythians preceding them, agreeing with the Gothic component in the multi-ethnic mix of the Chernyakhiv culture.

The higher proportion Near Eastern and (according to CP/NNLS) lower proportion of eastern ancestry in the Chernyakhiv culture samples were mirrored by f4 analyses where Chern showed lower affinity to Han (Z score –3.097) and EHG (Z score –3.643) than Ukrainian Scythian and Bronze Age samples, respectively, as well as higher Near Eastern (Levant_N and Anatolia_N) affinity than Ukrainian Scythians (Z scores 4.696 and 3.933, respectively). It is plausible to assume that this excess Near Eastern ancestry in Chern is related to European populations whose Near Eastern proportion has exceeded that in the steppe populations since the Neolithic expansion of early farmers. While the Chernyakhiv culture was likely ethnically heterogeneous, the three samples in our Chern group appear to represent its Gothic component.

chernyakhov-goths-uralic-clines
PCA obtained by projecting the ancient samples of this study together with published Scythian/Sarmatian and related samples onto a plot based on 537,802 autosomal SNPs in 1,422 modern Eurasians. To improve readability, the modern populations have been plotted as population medians (after outlier removal). Image modified from the paper, including Sredni Stog, Corded Ware/Uralic (with Srubna outliers) and Chernyakhov clusters.Notice the two new Late Yamna and Catacomb samples from Ukraine clustering with other published samples, despite being from the same region as Sredni Stog individuals.

Early Slavs of hg. I2-L621

A post in Anthrogenica shows some subclades of the varied haplogroups that are expected from medieval Poland:

KO_55, Kowalewko (100-300 AD), I1a3a1a1-Y6626
KO_45, Kowalewko (100-300 AD), I2a2a1b2a-L801
KO_22, Kowalewko (100-300 AD), G2a2b-L30
KO_57, Kowalewko (100-300 AD), G2a2b-L30

ME_7, Markowice (1000-1200 AD), I1a2a2a5-Y5384
NA_13, Niemcza, (900-1000 AD), I2a1b2-L621
NA_18, Niemcza, (900-1000 AD), J2a1a-L26

Just because of these samples among Early Slavs, and looking again more carefully at the modern distribution of I2a-L621 subclades, I think now I was wrong in assuming that I2a-L621 in early Hungarian Conquerors would mean they would appear around the Urals as a lineage integrated in Eastern Corded Ware groups. It seems rather a haplogroup with an origin in Central Europe. Whether it was part of a Baltic community that expanded south, or was incorporated during the expansions to the south is unclear. Like hg. E-V13, it doesn’t seem to have been incorporated precisely along the Danube, but closer to the north-east Carpathians.

Especially interesting is the finding of I2a-L621 among Early Slavs from Silesia, a zone of close interaction among early West Slavs. From Curta (2019):

On Common Slavs

In Poland, settlement discontinuity was postulated, to make room for the new, Prague culture introduced gradually from the southeast (from neighboring Ukraine). However, there is increasing evidence of 6th-century settlements in Lower Silesia (western Poland and the lands along the Middle Oder) that have nothing to do with the Prague culture. Nor is it clear how and when did the Prague culture spread over the entire territory of Poland.

On Great Moravia

Svatopluk’s remarkably strong position was immediately recognized by Pope John VIII, who ordered the immediate release of Methodius from his monastic prison in order to place him in 873 under Svatopluk’s protection. One year later (874), Louis the German himself was forced to recognize Svatopluk’s independence through the peace of Forchheim. By that time, the power of Svatopluk had extended into the upper Vistula Basin, over Bohemia, the lands between the Saale and the Elbe rivers, as well as the northern and northeastern parts of the Carpathian Basin.* The Czech prince Bořivoj, a member of the Přemyslid family which would unify and rule Bohemia in the following century, is believed to have been baptized in 874 by Methodius in Moravia together with his wife Ludmila (St. Wenceslas’s grandmother).

*Brather, Archäologie, p. 71. The expansion into the region of the Upper Vistula (Little Poland) results from one of St. Methodius’ prophecies, for which see the Life of Methodius 11, p. 72; Poleski, “Contacts between the Great Moravian empire and the tribes”; Poleski, “Contacts between the tribes in the basins.” Despite an early recognition of the Moravian influences on the material culture in 9th-century southern Poland and Silesia (e.g., Dostál, “Das Vordringen”), the question of Svatopluk’s expansion has triggered in the 1990s a fierce debate among Polish archaeologists. See Wachowski, “Problem”; Abłamowicz, “Górny Śląsk”; Wachowski, “Północny zasięg ekspansji”; Szydłowski, “Czy ślad”; Jaworski, “Elemente.”

On Piast Poland

Mieszko agreed to marry Oda, the daughter of the margrave of the North March, for his first wife had died in 977. The marriage signaled a change in the relations with the Empire, for Mieszko sent troops to help Otto II against the Slavic rebels of 983. He also attacked Bohemia and incorporated Silesia and Lesser Poland into the Piast realm, which prompted Bohemians to ally themselves with the Slavic rebels against whom Emperor Otto was now fighting. By 980, therefore, Mieszko was part of a broader configuration of power, and his political stature was recognized in Scandinavia as well. His daughter, Swietoslawa married first Erik Segersäll of Sweden (ca. 970–ca. 995) and then Sweyn Forkbeard of Denmark (986–1014).26 In the early 990s, together with his wife and children, Mieszko offered his state (called “civitas Schinesghe,” the state of Gniezno) to the pope as a fief, as attested by a unique document known as Dagome iudex and preserved in a late 11th-century summary. The document describes the inner boundaries of the state and peripheral provinces, as if Gniezno were a civitas (city) in Italy, with its surrounding territory. Regional centers, however, did indeed come into being shortly before AD 1000 in Lesser Poland (Cracow, Sandomierz), Pomerania (Gdańsk), and Silesia (Wrocław). Such regional centers came to be distinguished from other strongholds by virtue of the presence within their walls of some of the earliest churches built in stone. Mieszko got his own, probably missionary bishop.

In light of this recent find, which complements the Early Slav of the High Middle Ages from Sunghir (ca. AD 1100-1200), probably from the Vladimir-Suzdalian Rus’, we can assume now less speculatively that I2a-CTS10228 most likely expanded with Common Slavs, because alternative explanations for its emergence in the Carpathian Basin, among Early West Slavs, and among Early East Slavs within this short period of time requires too many unacceptable assumptions.

dinaric-i2a-distribution
Modern distribution of “Dinaric” I2a. Modified from Balanovsky et al. (2008)

Hungarian Conquerors

Knowing that R1a-Z280 was an Eastern Corded Ware lineage, found from Baltic Finns to Finno-Ugric populations of the Trans-Urals, we can probably assign expanding Magyars to at least R1a-Z280, R1a-Z93, and N1c-L392 (xB197) lineages.

From Curta (2019):

Earlier Latin sources, especially those of the first half of the 10th century, refer to Magyars as Huns or Avars. They most likely called themselves Magyars, a word indicating that the language they spoke was not Turkic, but Finno-Ugrian, related to a number of languages spoken in Western Siberia and the southern Ural region. The modern word—Hungarian—derives from the Slavic word for those people, U(n)gri, which is another indication of Ugric roots. This has encouraged the search for the origin of the Hungarian people in the lands to the east from the Ural Mountains, in western Siberia, where the Hungarian language is believed to have emerged between 1000 and 500 BC.

In looking for the Magyar primordial homeland, they draw comparisons with the assemblages found in Hungary that have been dated to the 10th century and attributed to the Magyars. Some of those comparisons had extraordinary results. For example, the excavation of the burial mound cemetery recently discovered near Lake Uelgi, in the Cheliabinsk region of Russia, has produced rosette-shaped harness mounts and silver objects ornamented with palmette and floral designs arranged in reticulated patterns, which are very similar to those of Hungary. But Uelgi is not dated to prehistory, and many finds from that site coincided in time with those found in burial assemblages in Hungary. In other words, although there can be no doubt about the relations between Uelgi and the sites in Hungary attributed to the first generations of Magyars, those relations indicate a migration directly from the Trans-Ural lands, and not gradually, with several other stops in the forest-steppe and steppe zones of Eastern Europe. In the lands west of the Ural Mountains, the Magyars are now associated with the Kushnarenkovo (6th to 8th century) and Karaiakupovo (8th to 10th century) cultures, and with such burial sites as Sterlitamak (near Ufa, Bashkortostan) and Bol’shie Tigany (near Chistopol, Tatarstan).14 However, the same problem with chronology makes it difficult to draw the model of a migration from the lands along the Middle Volga. Many parallels for the so typically Magyar sabretache plates found in Hungary are from that region. They have traditionally been dated to the 9th century, but more recent studies point to the coincidence in time between specimens found in Eastern Europe and those from Hungary.

Adding J2a and I1a samples to the Early Slavic stock, based on medieval samples from Poland – with G2a and E-V13 lineages probably shared with Goths from Wielbark/Chernyakhov, or becoming acculturated in the Carpathian Basin – one is left to wonder which of these lineages actually took part in Common Slavic migrations/acculturation events, whenever and wherever those actually happened.

I have tentatively re-assigned lineages of Hungarian conquerors according to their likely origins in a simplistic way – similar to how the paper classifies them – , now (I think) less speculatively, assuming that Early Slavs likely formed eventually part of them:

hungarian-conquerors-y-dna-slavs
Image modified from the paper, with drawn red square around lineages of likely East Slavic origin, and blue squares around R1a-Z93, R1a-Z283, N1a-Z1936, and N1a-M2004 samples, of likely Ugric origin Y-Hg-s determined from 46 males grouped according to sample age, cemetery and Hg. Hg designations are given according to ISOGG Tree 2019. Grey shading designate distinguished individuals with rich grave goods, color shadings denote geographic origin of Hg-s according to Fig. 1. For samples K3/1 and K3/3 the innermost Hg defining marker U106* was not covered, but had been determined previously.

NOTE. The ancestral origin of lineages is meaningless for an ethnolinguistic identification. The only reasonable assumption is that all the individuals sampled formed part of the Magyar polity, shared Magyar culture, and likely spoke Hungarian, unless there is a clear reason to deny this: which I guess should include at least a clearly ‘foreign’ ancestry (showing a distant cluster compared to the group formed by all other samples), ‘foreign’ isotopic data (showing that he was born and/or raised outside of the Carpathian Basin), and particularly ‘foreign’ cultural assemblage of the burial, if one really wants to risk assuming that the individual didn’t speak Hungarian as his mother tongue.

“Dinaric” or Slavic I2a?

I don’t like the use of “Dinaric I2a”, because it is reminiscent of the use of “Iberian R1b-DF27”, or “Germanic R1b-U106”, when ancient DNA has shown that this terminology is most often wrong, and turns out to be misleading. As misleading as “Slavic R1a”. Recently, a Spanish reader wrote me emails wondering how could I possibly say that R1b-DF27 came from Central Europe, because modern distribution maps (see below) made it evident that the haplogroup expanded from Iberia…

DF27-iberia-france-m167
Contour maps of the derived allele frequencies of the SNPs analyzed in Solé-Morata et al. (2017).

The obvious answer is, these maps show modern distributions, not ancient ones. In the case of R1b-DF27, different Iberian lineages are not even related to the same expansion. At least R1b-M167/SRY2627 lineages seem to have expanded from Central Europe into Iberia much more recently than other DF27 subclades associated with Bell Beakers. What’s more, if R1b-M167/SRY2627 appear densest in north-east Spain it is not because of the impact of Celts or Iberians before the arrival of Romans, but because of the impact of medieval expansions during the Reconquista from northern kingdoms expanding south in the Middle Ages:

iberian-medieval-kingdoms-expansion-population-genomics
Genetic differentiation and the footprints of historical migrations in the Iberian Peninsula. Image modified from Bycroft et al. (2018).

Similarly, the term “Dinaric I2a”, based on the higher density in the Western Balkans, is misleading because it is probably the result of later bottlenecks. Just like the density of different R1a subclades among Modern Slavs is most likely the result of acculturation of different groups, especially to the east and north-east, where language shift is known to have happened in historical times, with the cradle of Russians in particular being a Finno-Volgaic hotspot, later expanding with hg. R1a-Z280 and N1c-L392 lineages.

Now, one may think that maybe Slavs expanded with ALL of these different lineages. Since we are talking about late Iron Age / medieval expansions, there might be confederations of different peoples expanding with a single lingua franca… But no, not really. Not likely in linguistics, not likely in archaeology, and apparently not in population genomics, either.

How many ancient peoples from the Iron Age and Early Middle Ages expanded with so many different lineages? We see bottlenecks in expansions even in recent times: say, in Visigoths under E-V13 (probably recently incorporated during their migrations); in Moors (mostly Berbers) with E-M81 and J; in medieval Iberians under different DF27 bottlenecks during the Reconquista (including huge bottlenecks among Basques); similarly, huge bottlenecks are found in Finnic expansions under N1c…How likely is it that Proto-Slavs (and Common Slavs) expanded with all those attested lineages to date among Early Slavs (E-V13, I2a-L621, R1a-M458, I1, J2a) AND also with other R1a subclades prevalent today, but almost absent in sampled Early Slavs?

To sum up, I am not so sure anymore about the possibility of simplistically assigning R1a-M458 to expanding Common Slavs. R1a-M458 may well have been the prevalent R1a subclade in Central Europe among early Balto-Slavic – and possibly also neighbouring Northern Indo-European-speaking – peoples (let’s see what subclades Tollense and Unetice samples bring), but it is more and more likely that most of the density we see in modern R1a-M458 distribution maps is actually the effect of medieval bottlenecks of West Slavs, similar to the case of Iberia.

r1a-m458-underhill-2015
Modern distribution of R1a-M458, after Underhill et al. (2015).

Related

Baltic Finns in the Bronze Age, of hg. R1a-Z283 and Corded Ware ancestry

estonian-bronze-age-dna

Open access The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East, by Saag et al. Current Biology (2019).

Interesting excerpts:

In this study, we present new genomic data from Estonian Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). The cultural background of stone-cist graves indicates strong connections both to the west and the east [20, 21]. The Iron Age (IA) tarands have been proposed to mirror “houses of the dead” found among Uralic peoples of the Volga-Kama region [22].

(…) The 33 individuals included 15 from EstBA, 6 from EstIA, 5 from Pre-Roman to Roman Iron Age Ingria (500 BC–450 AD) (IngIA), and 7 from Middle Age Estonia (1200–1600 AD) (EstMA) and yielded endogenous DNA ∼4%–88%, average genomic coverages ∼0.017–0.734×, and contamination estimates <4% (Table S1). We analyzed the data in the context of modern and other ancient individuals, including from Neolithic Estonia [13].

estonian-y-dna-bronze-iron-age
Archaeological Information, Genetic Sex, mtDNA and Y Chromosome Haplogroups, and Average Coverage of the Individuals of This Study. Modified from the paper to mark distinct Y-DNA haplogroups in the LBA and IA.

We identified chrY hgs for 30 male individuals (Tables 1 and S2; STAR Methods). All 16 successfully haplogrouped EstBA males belonged to hg R1a, showing no change from the CWC period, when this was also the only chrY lineage detected in the Eastern Baltic [11, 13, 30, 31]. Three EstIA and two IngIA individuals also belonged to hg R1a, but three EstIA males belonged to hg N3a, the earliest so far observed in the Eastern Baltic. Three EstMA individuals belonged to hg N3a, two to hg R1a, and one to hg J2b. ChrY lineages found in the Baltic Sea region before the CWC belong to hgs I, R1b, R1a5, and Q [10, 11, 12, 13, 17, 32]. Thus, it appears that these lineages were substantially replaced in the Eastern Baltic by hg R1a [10, 11, 12, 13], most likely through steppe migrations from the east [30, 31]. (…) Our results enable us to conclude that, although the expansion time for R1a1 and N3a3′5 in Eastern Europe is similar [25], hg N3a likely reached Estonia or at least became comparably frequent to modern Estonia [1] only during the BA-IA transition.

A clear shift toward West Eurasian hunter-gatherers is visible between European LN and BA (including Baltic CWC) and EstBA individuals, the latter clustering together with Latvian and Lithuanian BA individuals [11]. EstIA, IngIA, and EstMA individuals project between BA individuals and modern Estonians, partially overlapping with both.

(…) EstBA individuals are clearly distinguishable from Estonian CWC individuals as the former have more of the blue component most frequent in WHGs and less of the brown and yellow components maximized in Caucasus hunter-gatherers and modern Khanty, respectively. The individuals of EstBA, EstIA, IngIA, EstMA, and modern Estonia are quite similar to each other on average, indicating that the relatively high proportion of WHG ancestry in modern Eastern Baltic populations compared to other present-day Europeans [15] traces back to the BA.

estonian-pca-published
Detail of the PCA, modified from the paper to label populations. Estonian Bronze Age and Iron Age samples cluster close to Early Corded Ware from the Baltic.. Principal-component analysis results of modern West Eurasians with ancient individuals projected onto the first two components (PC1 and PC2). BA, Bronze Age; EF, early farmers; HG, hunter-gatherers; IA, Iron Age; IMA, Iron/Middle Ages; LN, Late Neolithic; LNBA, Late Neolithic/Bronze Age; MA, Middle Ages

When comparing Estonian CWC and EstBA using autosomal outgroup f3 and Patterson’s D statistics (Table S3), the latter is more similar to other Baltic BA populations, to Baltic IA and Middle Age (MA) populations, and also to populations similar to WHGs and Scandinavian hunter-gatherers (SHGs), but not to Estonian CCC (Figures 2A and S2A; Data S1). The increase in WHG or SHG ancestry could be connected to western influences seen in material culture [20, 21] and facilitated by a decline in local population after the CCC-CWC period [20]. A slight trend of bigger similarity of Estonian CWC to forest or steppe zone populations and of EstBA to European early farmer populations can also be seen.

(…) When comparing to modern populations, Estonian CWC is slightly more similar to Caucasus individuals but EstBA to Baltic populations and Finnic speakers (Figure 2B; Data S1). Outgroup f3 and D statistics do not reveal apparent differences when comparing EstBA to EstIA, EstIA to IngIA, and EstIA to EstMA (Data S1).

estonian-ba-ia-ancestry
qpAdm results. Error bars indicate one SE. Central MN, Central European Middle Neolithic; EstBA, Estonian Bronze Age; EstIA, Estonian Iron Age; IngIA, Ingrian Iron Age; EstMA, Estonian Middle Ages; WHG, western hunter-gatherers.

These results highlight how uniparental and autosomal data can lead to different demographic inferences—the genetic change between CWC and BA not seen in uniparental lineages is clear in autosomal data and the appearance of chrY hg N in the IA is not matched by a clear shift in autosomal profiles.

EstBA individuals have no Nganasan-related ancestry and EstIA, IngIA, and EstMA individuals on average have 2% or 4% (Figure 3; Data S1). The differentiation remains when using BA or IA Fennoscandian populations [26] instead of Nganasans (Data S1). Notably, the proportion of Nganasan-related ancestry varies between 0% and 12% among sampled EstIA, IngIA, and EstMA individuals (Data S1), which may suggest its relatively recent admixture into the target population. Moreover, two individuals from Kunda (0LS10 and V10) have the highest proportions of Nganasan ancestry among EstIA (6% and 8%), one of them has chrY hg N3a, and isotopic analysis suggests neither individual being born in Kunda [34].

About these two males from Tarand-graves, ‘foreign’ to Kunda:

0LS10: Male from tarand III (burial 9; TÜ 1325: L777), age 17–25 years [34]. He had a fragment of a sheep/goat bone and ceramics as grave goods. This burial has two radiocarbon dates: 2430 ± 35 BP (Poz-10801; 760–400 cal BC) and 2530 ± 41 BP (UBA-26114; 800–530 cal BC) [34]. According to the isotopic analysis, the person was not born in the vicinity of Kunda; his place of birth is still unknown (but south-western Finland and Sweden are excluded) [34]. Sampled tooth r P1.

V10: Male from tarand XI (burial 24; TÜ 1325: L1925), age 25–35 years [34], date 2484 ± 40 BP (UBA-26115; 790–430 cal BC) [34]. He had a few potsherds near the skull. Likewise, this person was not locally born [34]. Sampled tooth l P1.

estonia-bronze-iron-age-steppe-siberian
Autosomal Analyses’ Results for Gyvakarai1 as the closest available Corded Ware source for Balto-Finnic populations.

The paper shows thus:

  • Major continuity of ancestry from Corded Ware to modern Estonians, with only slight changes in different periods. In fact, one of the best fits for the Late Bronze Age ancestry is Gyvakarai1, one of the Corded Ware “outliers” described as “closer to Yamna”, which I already said may be closer to Sredni Stog/EHG populations instead. Another interesting take is that the change from Bronze Age to Iron Age corresponds to an increase in Baltic Corded Ware-related ancestry, rather than being driven by Siberian ancestry.
  • pca-mittnik-gyvakarai
    File modified by me from Mittnik et al. (2018) to include the approximate position of the most common ancestral components, and an identification of potential outliers. Zoomed-in version of the European Late Neolithic and Bronze Age samples. “Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). From Mittnik et al. (2018).
  • A Volosovo-related migration of hg. N1c with Netted Ware into the area seems to be discarded, based on the full replacement of paternal lines and continuity of R1a-Z283. It is only during the Tarand-grave period when a system of chiefdoms (spread from Ananyino/Akozino) brings haplogroup N1c to the Gulf of Finland. During the Iron Age, the proportion of paternal lineages is still clearly in favour of R1a (50% in the coast, 100% in Ostrobothnia), which indicates a gradual replacement led by elites, likely because of the incorporation of Akozino warrior-traders spreading all over the Baltic, bringing the described shared Mordvinic traits in Fennic.
  • finno-ugric-haplogroup-n
    Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).
  • The arrival of Akozino warrior-traders (bringing N1c and R1a lineages) was probably linked to this minimal “Nganasan-like” ancestry of some samples in the transition to the Iron Age. This arrival is supported by samples 0LS10 (the earliest hg. N1c) and V10 (of hg. R1a), both dated to ca. 800-400 BC, with V10 showing the highest “Nganasan-like” ancestry with 4.8%, both of them neighbouring samples showing 0%. This variable admixture among local and foreign paternal lineages might support the described social system of family alliances with intermarriages. In fact, a medieval sample, 0LS03_1 (hg. R1a) also shows a recent “Nganasan-like” ancestry, which probably points to the integration of different Arctic-related ancestry components among Modern Estonians, in this case related to Finnish expansions and thus integration of Levänluhta-related ancestry, as per the supplementary data.
  • NOTE. Such minimal proportions of “Nganasan-like” ancestry evidence the process of admixture of Volga Finns in Akozino territory through their close interactions with Permians of Ananyino, who in turn acquired this Palaeo-Arctic admixture most likely during the expansion of the linguistic community to hunter-gatherer territories, to the north of the Cis-Urals. This process of stepped infiltration and expansion without language change is not dissimilar to the one seen among Indo-Iranians and Balto-Slavs of hg. R1b, or Vasconic speakers of hg. I2a, although in the case of Baltic Finns of hg. R1a the process of infiltration and expansion of hg. N1c is much less dramatic, with no radical replacement anywhere before the huge bottlenecks observable in Finns.

  • The expansion of haplogroup N1c among Finnic populations, as we are going to see in samples from the Middle Ages such as Luistari, is the consequence of late founder effects after huge bottlenecks expected based on the analysis of modern populations. The expansion of N1c-VL29 is different in origin from that of N1c-Z1936 among Samic (later integrated into Finnish populations), most likely from the east and originally associated with Lovozero Ware.
haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders. Map from Ilumäe et al. (2016).

In spite of all this, the conclusion of the paper is (surprise!) that Siberian ancestry and hg. N heralded the arrival of Finnic to the Gulf of Finland in the Iron Age… However, this conclusion is supposedly* supported, not by their previous papers, but by a recent phylogenetic study by Honkola et al. (2013), which doesn’t actually argue for such a late ‘arrival’: it argues for the split of Balto-Finnic around 1500 BC.

NOTE. I say ‘supposedly’ because Kristiina Tambets, for example, has been following the link of Uralic with haplogroup N since the 2000s, so this is not some conclusion they just happened to misread from some random paper they Googled. In those initial assessments, she argued that the “ancient homeland” of the Tat C mutation suggested that Finno-Ugrians were in Fennoscandia before Indo-Europeans. Apparently, since haplogroup N appears later and from the east, it is now more important to follow this haplogroup than what is established in archaeology and linguistics.

Even in the referred paper, this split is considered an in situ development, since the phylogenetic study takes the information – among others – 1) from Parpola and Carpelan, who consider Netted Ware, a culture derived from Fatyanovo/Abashevo and Volosovo, as the culprit of the Finno-Ugric expansion; and 2) from Kallio (2006), who clearly states that Proto-Balto-Finnic (like Proto-Finno-Samic) was spoken around the Gulf of Finland during the Bronze Age. Both of them set the terminus ante quem of the language presence in the Baltic ca. 1900 BC.

Anyways, as a consequence of geneticists keeping these untenable pre-ancient DNA haplogroup-based arguments today, I expect to see this “Finnic” language expansion also described for the Western Baltic, Scandinavia or northern Europe, when this same proportion of hg. N1c and “Nganasan” ancestry is observed in Iron Age samples around the Baltic Sea. The nativist trends that this domination of “Finns” all over Northern Europe 2,500 years ago will create will be even more fun to read than the current ones…

EDIT (10 May 2019) How I see the reaction of many to ancient DNA, in keeping their old theories:

Related

Złota a GAC-CWC transitional group…but not the origin of Corded Ware peoples

koszyce-gac-zlota-cwc

Open access Unraveling ancestry, kinship, and violence in a Late Neolithic mass grave, by Schroeder et al. PNAS (2019).

Interesting excerpts of the paper and supplementary materials, about the Złota group variant of Globular Amphora (emphasis mine):

A special case is the so-called Złota group, which emerged around 2,900 BCE in the northern part of the Małopolska Upland and existed until 2,600-2,500 BCE. Originally defined as a separate archaeological “culture” (15), this group is mainly defined by the rather local introduction of a distinct form of burial in the area mentioned. Distinct Złota settlements have not yet been identified. Nonetheless, because of the character of its burial practices and material culture, which both retain many elements of the GAC and yet point forward to the Corded Ware tradition, and because of its geographical location, the Złota group has attracted significant archaeological attention (15, 16).

The Złota group buried their dead in a new, distinct type of funerary structure; so-called niche graves (also called catacomb graves). These structures featured an entrance shaft or pit and, below that, a more or less extensive niche, sometimes connected to the entrance area by a narrow corridor. Local limestone was used to seal off the entrance shaft and to pave the floor of the niche, on which the dead were usually placed along with grave goods. This specific and relatively sophisticated form of burial probably reflects contacts between the northern Małopolska Upland and the steppe and forest-steppe communities further to the east, who also buried their dead in a form of catacomb graves. Individual cases of the use of ochre and of deformation of skulls in Złota burials provide further indications of such a connection (15). At the same time, the Złota niche grave practice also retains central elements of the GAC funerary tradition, such as the frequent practice of multiple burials in one grave, often entailing redeposition and violation of the anatomical order of corpses, and thus differs from the catacomb grave customs found on the steppes which are strongly dominated by single graves. Nonetheless, at Złota group cemeteries single burial graves appear, and even in multiple burial graves the identity of each individual is increasingly emphasized, e.g. by careful deposition of the body and through the personal nature of grave goods (16).

globular-amphorae-corded-ware-zlota-amphorae
Correspondence analysis of amphorae from the Złota-graveyards reveals that there is no typological break between Globular Amphorae and Corded Ware Amphorae, including ‘Strichbündelamphorae’ (after Furholt 2008)

Just like its burial practices, the material culture and grave goods of the Złota group combine elements of the GAC, such as amber ornaments and central parts of the ceramic inventory, with elements also found in the Corded Ware tradition, such as copper ornaments, stone shaft-hole axes, bone and shell ornaments, and other stylistic features of the ceramic inventory. In particular, Złota group ceramic styles have been seen as a clear transitional phenomenon between classical GAC styles and the subsequent Corded Ware ceramics, probably playing a key role in the development of the typical cord decoration patterns that came to define the latter (17).

As briefly summarized above, the Złota group displays a distinct funerary tradition and combination of material culture traits, which give the clear impression of a cultural “transitional situation”. While the group also appears to have had long-distance contacts directed elsewhere (e.g. to Baden communities to the south), it is the combination of Globular Amphora traits, on the one hand, and traits found among late Yamnaya or Catacomb Grave groups to the east as well as the closely related Corded Ware groups that emerged around 2,800 BCE, on the other hand, that is such a striking feature of the Złota group and which makes it interesting when attempting to understand cultural and demographic dynamics in Central and Eastern Europe during the early 3rd millennium BCE.

catacomb-grave-ksiaznice
Catacomb grave no. 2a/06 from Książnice, Złota culture (acc. to Wilk 2013). Image from Włodarczak (2017)

Książnice (site 2, grave 3ZC), Świętokrzyskie province. This burial, a so-called niche grave of the Złota type (with a vertical entrance shaft and perpendicularly situated niche), was excavated in 2006 and contained the remains of 8 individuals, osteologically identified as three adult females and five children, positioned on limestone pavement in the niche part of the grave. Radiocarbon dating of the human remains indicates that the grave dates to 2900-2630 BCE, 95.4% probability (Dataset S1). The grave had an oval entrance shaft with a diameter of 60 cm and depth of 130 cm; the depth of the niche reached to 170 cm (both measured from the modern surface), and it also contained a few animal bones, a few flint artefacts and four ceramic vessels typical of the Złota group. Książnice is located in the western part of the Małopolska Upland, which only has a few Złota group sites but a stronger presence of other, contemporary groups (including variants of the Baden culture).

Wilczyce (site 90, grave 10), Świętokrzyskie province. A rescue excavation in 2001 uncovered a niche grave of the Złota type, which had a round entrance shaft measuring 90 cm in diameter. The grave was some 60-65 cm deep below the modern surface and the bottom of the niche was paved with thin limestone plates, on which remains of three individuals had been placed; two adults, one female and one male, and one child. Four ceramic vessels of Złota group type were deposited in the niche along with the bodies. Wilczyce is located in the Sandomierz Upland, an area with substantial presence of both the Globular Amphora culture and Złota group, as well as the Corded Ware culture from 2800 BCE.

zlota-gac-cwc
Genetic affinities of the Koszyce individuals and other GAC groups (here including Złota) analyzed in this study. (A) Principal component analysis of previously published and newly sequenced ancient individuals. Ancient genomes were projected onto modern reference populations, shown in gray. (B) Ancestry proportions based on supervised ADMIXTURE analysis (K = 3), specifying Western hunter-gatherers, Anatolian Neolithic farmers, and early Bronze Age steppe populations as ancestral source populations. LP, Late Paleolithic; M, Mesolithic; EN, Early Neolithic; MN, Middle Neolithic; LN, Late Neolithic; EBA, Early Bronze Age; PWC, Pitted Ware culture; TRB, Trichterbecherkultur/Funnelbeaker culture; LBK, Linearbandkeramik/Linear Pottery culture; GAC, Globular Amphora culture; Złota, Złota culture. Image modified to outline in red GAC and Złota groups.

To further investigate the ancestry of the Globular Amphora individuals, we performed a supervised ADMIXTURE (6) analysis, specifying typical western European hunter-gatherers (Loschbour), early Neolithic Anatolian farmers (Barcın), and early Bronze Age steppe populations (Yamnaya) as ancestral source populations (Fig. 2B). The results indicate that the Globular Amphora/Złota group individuals harbor ca. 30% western hunter-gatherer and 70% Neolithic farmer ancestry, but lack steppe ancestry. To formally test different admixture models and estimate mixture proportions, we then used qpAdm (7) and find that the Polish Globular Amphora/Złota group individuals can be modeled as a mix of western European hunter-gatherer (17%) and Anatolian Neolithic farmer (83%) ancestry (SI Appendix, Table S2), mirroring the results of previous studies.

zlota-steppe-ancestry-cwc
Table S2. qpADM results. The ancestry of most Globular Amphora/Złota group individuals
can be modelled as a two-way mixture of Mesolithic western hunter-gatherers (WHG), and early Anatolian Neolithic farmers (Barcın). The five individuals from Książnice (Złota group) show evidence for additional gene flow, most likely from an eastern source.

The lack of a direct genetic connection of Corded Ware peoples with the Złota group despite their common “steppe-like traits” – shared with Yamna – reveals, once more, how the few “Yamna-like” traits of Corded Ware do not support a direct connection with Indo-Europeans, and are the result of the expansion of the so-called steppe package all over Europe, and particularly among cultures closely related to the Khvalynsk expansion, and later under the influence of expanding Yamna peoples.

The results from Książnice may support that early Corded Ware peoples were in close contact with GAC peoples in Lesser Poland during the complex period of GAC-Trypillia-CWC interactions, and especially close to the Złota group at the beginning of the 3rd millennium BC. Nevertheless, patrilineal clans of Złota apparently correspond to Globular Amphorae populations, with the only male sample available yet being within haplogroup I2a-L801, prevalent in GAC.

NOTE. The ADMIXTURE of Złota samples in common with GAC samples (and in contrast with the shared Sredni Stog – Corded Ware “steppe ancestry”) makes the possibility of R1a-M417 popping up in the Złota group from now on highly unlikely. If it happened, that would complicate further the available picture of unusually diverse patrilineal clans found among Uralic speakers expanding with early Corded Ware groups, in contrast with the strict patrilineal and patrilocal culture of Indo-Europeans as found in Repin, Yamna and Bell Beakers.

Once again the traditional links between groups hypothesized by archaeologists – like Gimbutas and Kristiansen in this case – are wrong, as is the still fashionable trend in descriptive archaeology, of supporting 1) wide cultural relationships in spite of clear-cut inter-cultural differences (and intra-cultural uniformity kept over long distances by genetically-related groups), 2) peaceful interactions among groups based on few common traits, and 3) regional population continuities despite cultural change. These generalized ideas made some propose a steppe language shared between Pontic-Caspian groups, most of which have been proven to be radically different in culture and genetics.

gimbutas-kurgan-indo-european
The background shading indicates the tree migratory waves proposed by Marija Gimbutas, and personally checked by her in 1995. Image from Tassi et al. (2017).

Furthermore, paternal lines show once again marked bottlenecks in expanding Neolithic cultures, supporting their relevance to follow the ethnolinguistic identity of different cultural groups. The steppe- or EHG-related ancestry (if it is in fact from early Corded Ware peoples) in Książnice was thus probably, as in the case of Trypillia, in the form of exogamy with females of neighbouring groups:

The presence of unrelated females and related males in the grave is interesting because it suggests that the community at Koszyce was organized along patrilineal lines of descent, adding to the mounting evidence that this was the dominant form of social organization among Late Neolithic communities in Central Europe. Usually, patrilineal forms of social organization go hand in hand with female exogamy (i.e., the practice of women marrying outside their social group). Indeed, several studies (11, 12) have shown that patrilocal residence patterns and female exogamy prevailed in several parts of Central Europe during the Late Neolithic. (…) the high diversity of mtDNA lineages, combined with the presence of only a single Y chromosome lineage, is certainly consistent with a patrilocal residence system.

funnelbeaker-trypillia-corded-ware
Map of territorial ranges of Funnel Beaker Culture (and its settlement concentrations in Lesser Poland), local Tripolyan groups and Corded Ware Culture settlements (■) at the turn of the 4th/3rd millennia BC.

Since ancient and modern Uralians show predominantly Corded Ware ancestry, and Proto-Uralic must have been in close contact with Proto-Indo-European for a very long time – given the different layers of influence that can be distinguished between them -, it follows as logical consequence that the North Pontic forest-steppes (immediately to the west of the PIE homeland in the Don-Volga-Ural steppes) is the most likely candidate for the expansion of Proto-Uralic, accompanying the spread of Sredni Stog ancestry and a bottleneck under R1a-M417 lineages.

The early TMRCAs in the 4th millennium BC for R1a-M417 and R1a-Z645 support this interpretation, like the R1a-M417 sample found in Sredni Stog. On the other hand, the resurgence of typical GAC-like ancestry in late Corded Ware groups, with GAC lineages showing late TMRCAs in the 3rd millennium BC, proves the disintegration of Corded Ware all over Europe (except in Textile Ceramics- and Abashevo-related groups) as the culture lost its cohesion and different local patrilineal clans used the opportunity to seize power – similar to how eventually I2a-L621 infiltrated eastern (Finno-Ugrian) groups.

Related

Yamna the likely source of modern horse domesticates; the closest lineage, from East Bell Beakers

Open access Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series, by Fages et al. Cell (2019).

Interesting excerpts (emphasis mine):

The earliest archaeological evidence of horse milking, harnessing, and corralling is found in the ∼5,500-year-old Botai culture of Central Asian steppes (Gaunitz et al., 2018, Outram et al., 2009; see Kosintsev and Kuznetsov, 2013 for discussion). Botai-like horses are, however, not the direct ancestors of modern domesticates but of Przewalski’s horses (Gaunitz et al., 2018). The genetic origin of modern domesticates thus remains contentious, with suggested candidates in the Pontic-Caspian steppes (Anthony, 2007), Anatolia (Arbuckle, 2012, Benecke, 2006), and Iberia (Uerpmann, 1990, Warmuth et al., 2011). Irrespective of the origins of domestication, the horse genome is known to have been reshaped significantly within the last ∼2,300 years (Librado et al., 2017, Wallner et al., 2017, Wutke et al., 2018). However, when and in which context(s) such changes occurred remains largely unknown.

To clarify the origins of domestic horses and reveal their subsequent transformation by past equestrian civilizations, we generated DNA data from 278 equine subfossils with ages mostly spanning the last six millennia (n = 265, 95%) (Figures 1A and 1B; Table S1; STAR Methods). Endogenous DNA content was compatible with economical sequencing of 87 new horse genomes to an average depth-of-coverage of 1.0- to 9.3-fold (median = 3.3-fold; Table S2). This more than doubles the number of ancient horse genomes hitherto characterized. With a total of 129 ancient genomes, 30 modern genomes, and new genome-scale data from 132 ancient individuals (0.01- to 0.9-fold, median = 0.08-fold), our dataset represents the largest genome-scale time series published for a non-human organism (Tables S2, S3, and S4; STAR Methods).

genetic-affinities-horse-domesticates-pca
Genetic Affinities.
(A)
Principal Component Analysis (PCA) of 159 ancient and modern horse genomes showing at least 1-fold average depth-of-coverage. The overall genetic structure is shown for the first three principal components, which summarize 11.6%, 10.4% and 8.2% of the total genetic variation, respectively. The two specimens MerzlyYar_Rus45_23789 and Dunaujvaros_Duk2_4077 discussed in the main text are highlighted. See also Figure S7 and Table S5 for further information.
(B) Visualization of the genetic affinities among individuals, as revealed by the struct-f4 algorithm and 878,475 f4 permutations. The f4 calculation was conditioned on nucleotide transversions present in all groups, with samples were grouped as in TreeMix analyses (Figure 3). In contrast to PCA, f4 permutations measure genetic drift along internal branches. They are thus more likely to reveal ancient population substructure.

Discovering Two Divergent and Extinct Lineages of Horses

Domestic and Przewalski’s horses are the only two extant horse lineages (Der Sarkissian et al., 2015). Another lineage was genetically identified from three bones dated to ∼43,000–5,000 years ago (Librado et al., 2015, Schubert et al., 2014a). It showed morphological affinities to an extinct horse species described as Equus lenensis (Boeskorov et al., 2018). We now find that this extinct lineage also extended to Southern Siberia, following the principal component analysis (PCA), phylogenetic, and f3-outgroup clustering of an ∼24,000-year-old specimen from the Tuva Republic within this group (Figures 3, 5A and S7A). This new specimen (MerzlyYar_Rus45_23789) carries an extremely divergent mtDNA only found in the New Siberian Islands some ∼33,200 years ago (Orlando et al., 2013) (Figure 6A; STAR Methods) and absent from the three bones previously sequenced. This suggests that a divergent ghost lineage of horses contributed to the genetic ancestry of MerzlyYar_Rus45_23789. However, both the timing and location of the genetic contact between E. lenensis and this ghost lineage remain unknown.

modern-horse-domesticates-przewalski-hungary
Population modeling of the demographic changes and admixture events in extant and extinct horse lineages. The two models presented show best fitting to the observed multi-dimensional SFS in momi2. The width of each branch scales with effective size variation, while colored dashed lines indicate admixture proportions and their directionality. The robustness of each model was inferred from 100 bootstrap pseudo-replicates. Time is shown in a linear scale up to 120,000 years ago and in a logarithmic scale above.

Modeling Demography and Admixture of Extinct and Extant Horse Lineages

Phylogenetic reconstructions without gene flow indicated that IBE differentiated prior to the divergence between DOM2 and Przewalski’s horses (Figure 3; STAR Methods). However, allowing for one migration edge in TreeMix suggested closer affinities with one single Hungarian DOM2 specimen from the 3rd mill. BCE (Dunaujvaros_Duk2_4077), with extensive genetic contribution (38.6%) from the branch ancestral to all horses (Figure S7B).This, and the extremely divergent IBE Y chromosome (Figure 6B), suggest that a divergent but yet unidentified ghost population could have contributed to the IBE genetic makeup.

Rejecting Iberian Contribution to Modern Domesticates

The genome sequences of four ∼4,800- to 3,900-year-old IBE specimens characterized here allowed us to clarify ongoing debates about the possible contribution of Iberia to horse domestication (Benecke, 2006, Uerpmann, 1990, Warmuth et al., 2011). Calculating the so-called fG ratio (Martin et al., 2015) provided a minimal boundary for the IBE contribution to DOM2 members (Cahill et al., 2013) (Figure 7A). The maximum of such estimate was found in the Hungarian Dunaujvaros_Duk2_4077 specimen (∼11.7%–12.2%), consistent with its TreeMix clustering with IBE when allowing for one migration edge (Figure S7B). This specimen was previously suggested to share ancestry with a yet-unidentified population (Gaunitz et al., 2018). Calculation of f4-statistics indicates that this population is not related to E. lenensis but to IBE (Figure 7B; STAR Methods). Therefore, IBE or horses closely related to IBE, contributed ancestry to animals found at an Early Bronze Age trade center in Hungary from the late 3rd mill. BCE. This could indicate that there was long-distance exchange of horses during the Bell Beaker phenomenon (Olalde et al., 2018). The fG minimal boundary for the IBE contribution into an Iron Age Spanish horse (ElsVilars_UE4618_2672) was still important (~9.6%–10.1%), suggesting that an IBE genetic influence persisted in Iberia until at least the 7th century BCE in a domestic context. However, fG estimates were more limited for almost all ancient and modern horses investigated (median = ~4.9%–5.4%; Figure 7A).

horse-lineages-domesticates-przewalski-dom2-botai
TreeMix Phylogenetic Relationships. The tree topology was inferred using a total of ∼16.8 million transversion sites and disregarding migration. The name of each sample provides the archaeological site as a prefix, and the age of the specimen as a suffix (years ago). Name suffixes (E) and (A) denote European and Asian ancient horses, respectively. See Table S5 for dataset information. Image modified to include the likely ancestor of domesticates in a red circle, represented by Yamna, the most likely direct ancestor of the Dunaujvarus specimen.

Iron Age horses

Y chromosome nucleotide diversity (π) decreased steadily in both continents during the last ∼2,000 years but dropped to present-day levels only after 850–1,350 CE (Figures 2B and S2E; STAR Methods). This is consistent with the dominance of an ∼1,000- to 700-year-old oriental haplogroup in most modern studs (Felkel et al., 2018, Wallner et al., 2017). Our data also indicate that the growing influence of specific stallion lines post-Renaissance (Wallner et al., 2017) was responsible for as much as a 3.8- to 10.0-fold drop in Y chromosome diversity.

We then calculated Y chromosome π estimates within past cultures represented by a minimum of three males to clarify the historical contexts that most impacted Y chromosome diversity. This confirmed the temporal trajectory observed above as Byzantine horses (287–861 CE) and horses from the Great Mongolian Empire (1,206–1,368 CE) showed limited yet larger-than-modern diversity. Bronze Age Deer Stone horses from Mongolia, medieval Aukštaičiai horses from Lithuania (C9th–C10th [ninth through the tenth centuries of the Common Era]), and Iron Age Pazyryk Scythian horses showed similar diversity levels (0.000256–0.000267) (Figure 2A). However, diversity was larger in La Tène, Roman, and Gallo-Roman horses, where Y-to-autosomal π ratios were close to 0.25. This contrasts to modern horses, where marked selection of specific patrilines drives Y-to-autosomal π ratios substantially below 0.25 (0.0193–0.0396) (Figure 2A). The close-to-0.25 Y-to-autosomal π ratios found in La Tène, Roman, and Gallo-Roman horses suggest breeding strategies involving an even reproductive success among stallions or equally biased reproductive success in both sexes (Wilson Sayres et al., 2014).

Lineage is used in this paper, as in many others in genetics, as defined by a specific ancestry. I keep that nomenclature below. It should not be confused with the “lineages” or “lines” referring to Y-chromosome (or mtDNA) haplogroups.

Supporting the “archaic” nature of the Hungarian BBC horses expanding from the Pontic-Caspian steppes are:

  • Among Y-chromosome lines, the common group formed by Botai-Borly4 (closely related to DOM2), Scythian horses from Aldy Bel (Arzhani), Iron Age horses from Estonia (Ridala), horses from the Xiongnu culture (Uushgiin Uvur), and Roman horses from Autricum (Chartres).
  • Among mtDNA lines, the common group formed by Botai samples, LebyazhinkaIV NB35, and different Eurasian domesticates, including many ancient Western European ones, which reveals a likely expansion of certain subclades east and west with the Repin culture.
  • (…) DOM2 contributed 22% to the ancestor of Przewalski’s horses ca. 9.47 kya, suggesting the Holocene optimum, rather than the Eneolithic Botai culture (∼5.5 kya), as a period of population contact. This pre-Botai introgression could explain the Y chromosome topology, where Botai horses were reported to carry two different segregating haplogroups: one occupied a basal position in the phylogeny while the other was closely related to DOM2. Multiple admixture pulses, however, are known to have occurred along the divergence of DOM2 and the Botai-Borly4 lineage, including 2.3% post-Borly4 contribution to DOM2, and a more recent 6.8% DOM2 intogression into Przewalski’s horses (Gaunitz et al., 2018). Model C2 parameters accommodate all these as a single admixture pulse, likely averaging the contributions of all these multiple events.

    horse-domesticate-y-dna-mtdna
    Tip labels are respectively composed of individual sample names, their reference number as well as their age (years ago, from 2017). Red, orange, light green, green, dark green and blue refer to modern horses, ancient DOM2, Botai horses, Borly4 horses, Przewalski’s horses and E. lenensis, respectively. Black refers to wild horses not yet identified to belong to any particular cluster in absence of sufficient genome-scale data. Clades composed of only Przewalski’s horses or ancient DOM2 horses were collapsed to increase readability.

    (A) Best maximum likelihood tree retracing the phylogenetic relationships between 270 mitochondrial genomes.

    B) Best Y chromosome maximum likelihood tree (GTRGAMMA substitution model) excluding outgroup. Node supports are indicated as fractions of 100 bootstrap pseudoreplicates. Bootstrap supports inferior to 90% are not shown. The root was placed on the tree midpoint. See also Table S5 for dataset information.

    Image modified from the paper, including a red square in archaic groups that contain the Hungarian sample, and a red circle around the most likely common ancestral stallion and mare from the Pontic-Caspian steppes.

    The paper cannot offer a detailed picture of ancient horse domestication, but it is yet another step in showing how Repin/Yamna is the most likely source of expansion of horse domesticates in Eurasia. Even more interestingly, Yamna settlers in Hungary probably expanded an ancient lineage of that horse at the same time as they spread with the Classical Bell Beaker culture. Remarkable parallels are thus found between:

    The expansion of an ancient line of horse domesticates related to Yamna Hungary/East Bell Beakers seems to be confirmed by the pre-Iberian sample from Vilars I, Els Vilars4618 2672 (ca. 700-550 BC), likely of Iberian Beaker descent, showing a lineage older than the Indo-Iranian ones, which later replaced most European lines.

    NOTE. For known contacts between Yamna and Proto-Beakers just before the expansion of East Bell Beakers, see a recent post on Vanguard Yamna groups.

    The findings of the paper confirm the expansion of the horse firstly (and mainly) through the steppe biome, mimicking the expansion of Proto-Indo-Europeans first, and then replaced gradually (or not so gradually) by lines brought to Europe during westward expansions of Bronze Age, Iron Age, and later specialized horse-riding steppe cultures. The expansion also correlates well with the known spread of animal traction and pastoralism before 2000 BC:

    animal-traction-europe
    Top image: Map with evidence of animal traction before ca. 2000 BC. Bottom image: frequency of finds of evidence for animal traction (orange), cylinder seals (purple) and potter’s wheels (green) in the 4th and 3rd millennium BC (query from the Digital Atlas of Innovations). The data points to an early peak in the expansion of this innovation at the turn of the 4th–3rd millennium BC, while direct evidence supports a radical increase from around the mid–3th millennium BC until the early 2nd millennium, coinciding with the expansion of East Bell Beakers and related European Early Bronze Age cultures. Data and image modified from Klimscha (2017).

    EDIT (3 MAY 2019): A recent reminder of these parallel developments by David Reich in Insights into language expansions from ancient DNA:

    • Yamna expansion to the west “with horses and wagons”, with a more homogeneous ancestry in modern Europeans due to later migrations from the east (and north):
    • “Descendants” of Yamna (once the culture was already “dead”), expanding to the east mainly with Corded Ware ancestry:

    Another recent open access paper on horse domestication is The horse Y chromosome as an informative marker for tracing sire lines, by Felkel et al. Scientific Reports (2019).

    Related

Uralic speakers formed clines of Corded Ware ancestry with WHG:ANE populations

steppe-forest-tundra-biomes-uralic

The preprint by Jeong et al. (2018) has been published: The genetic history of admixture across inner Eurasia Nature Ecol. Evol. (2019).

Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):

A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.

eurasian-clines-uralic-altaic
The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the northsouth cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals.

For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).

For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).

west-urals-finno-ugrians-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the 13 populations west of the Urals, we present a four-way admixture model, Nganasan+Srubnaya+WHG+LBK_EN, or its minimal adequate subset. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…

For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).

east-urals-ugric-samoyedic-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the four populations east of the Urals, we present three admixture models: Baikal_EBA+Srubnaya, Baikal_EBA+Srubnaya+AG3 and Nganasan+Srubnaya+AG3. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.

bronze-age-iron-age-karasuk-mezhovska-tagar-qpadm
Supplementary Table 9. QpAdm-based admixture modeling of Bronze and Iron Age populations of southern Siberia. For ancieint individuals associated with Karasuk and Tagar cultures, Nganasan+Srubnaya model is insufficient. For all five groups, adding AG3 as the third ancestry or substituting Nganasan with Baikal_EBA with higher ANE affinity provides an adequate model. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Models with p-value ≥ 0.05 are highlighted in bold face. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Lara M. Cassidy comments the results of the study in A steppe in the right direction (you can read it here):

Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).

Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.

eurasian-clines-uralic-turkic-mongol-altaic
The genomic formation of inner Eurasians. b–d, Depiction of the three main clines of ancestry identified among Inner Eurasians. Sources of admixture for each cline are represented using proxy ancient populations, both sampled and hypothesised, based on the study’s modelling results. The major eastern and western ancestries used to model each cline are shown in bold; the peripheral admixtures that gave rise to these are also shown. Additional contributions to subsections of each cline are marked with dashed lines. b, The northernmost cline, illustrating the legacy of WHG and ANE-related populations. c,d, The upper (c) and lower (d) steppe clines are shown, both of which have substantial eastern contributions related to modern Tungusic speakers. The authors propose these populations are themselves the result of an admixture between groups related to the Nganasan, whose ancestors potentially occupied a wider range, and hunter-gatherers (HGs) from the Amur River Basin. While the upper steppe cline in c can be described as a mixture between this eastern ancestry and western steppe herders, the current model for the southern steppe cline as shown in d is not adequate and is likely confounded by interactions with diverse bordering ancestries. Credit: Ecoregions 2017, Resolve https://ecoregions2017.appspot.com/

Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:

siberian-clines-uralic-altaic
PCA of ancient and modern Eurasian samples. Ancient Palaeo-Laplandic, Palaeosiberian, and Altai clines drawn, with modern populations labelled. See a version with higher resolution.

The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:

uralic-admixture-qpadm
Characterization of the Western and Eastern Eurasian source ancestries in inner Eurasian populations. [Modified from the paper, includes only Uralic populations]. a, Admixture f3 values are compared for different Eastern Eurasian (Mixe, Nganasan and Ulchi; green) and Western Eurasian references (Srubnaya and Chalcolithic Iranians (Iran_ChL); red). For each target group, darker shades mark more negative f3 values. b, Weights of donor populations in two sources characterizing the main admixture signal (date 1 and PC1) in the GLOBETROTTER analysis. We merged 167 donor populations into 12 groups (top right). Target populations were split into five groups (from top to bottom): Aleuts; the forest-tundra cline populations; the steppe-forest cline populations; the southern steppe cline populations; and ‘others’.

Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.

west-eurasian-east-eurasian-ancestry
Supplementary Fig. 4. ADMIXTURE results qualitatively support PCA-based grouping of inner Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”). Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”).

A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.

Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…

Related

Fulani from Cameroon show ancestry similar to Afroasiatic speakers from East Africa

sahel-region-fulani

Open access African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations, by Fan et al. Genome Biology (2019) 20:82.

Interesting excerpts (emphasis mine):

Introduction

To extend our knowledge of patterns of genomic diversity in Africa, we generated high coverage (> 30×) genome sequencing data from 43 geographically diverse Africans originating from 22 ethnic groups, representing a broad array of ethnic, linguistic, cultural, and geographic diversity (Additional file 1: Table S1). These include a number of populations of anthropological interest that have never previously been characterized for high-coverage genome sequence diversity such as Afroasiatic-speaking El Molo fishermen and Nilo-Saharan-speaking Ogiek hunter-gatherers (Kenya); Afroasiatic-speaking Aari, Agaw, and Amhara agro-pastoralists (Ethiopia); Niger-Congo-speaking Fulani pastoralists (Cameroon); Nilo-Saharan-speaking Kaba (Central African Republic, CAR); and Laka and Bulala (Chad) among others. We integrated this data with 49 whole genome sequences generated as part of the Simons Genome Diversity Project (SGDP) [14] (…)

afroasiatic-samples
Locations of samples included in this study. Each dot is an individual and the color indicates the language classification

Results and discussion

We found that the CRHG populations from central Africa, including the Mbuti from the Demographic Republic of Congo (DRC), Biaka from the CAR, and Baka, Bakola, and Bedzan from Cameroon, also form a basal lineage in the phylogeny. The other two hunter-gatherer populations, Hadza and Sandawe, living in Tanzania, group with populations from eastern Africa (Fig. 2). The two Nilo-Saharan-speaking populations, the Mursi from southern Ethiopia and the Dinka from southern Sudan, group into a single cluster, which is consistent with archeological data indicating that the migration of Nilo-Saharan populations to eastern Africa originated from a source population in southern Sudan in the last 3000 years [4, 23, 24, 25].

phylogenetic-relationship-africans
Phylogenetic relationship of 44 African and 32 west Eurasian populations determined by a neighbor joining analysis assuming no admixture. Here, the dots of each node represent bootstrap values and the color of each branch indicates language usage of each population. Human_AA human ancestral alleles

The Fulani people are traditionally nomadic pastoralists living across a broad geographic range spanning Sudan, the Sahel, Central, and Western Africa. The Fulani in our study, sampled from Cameroon, clustered with the Afroasiatic-speaking populations in East Africa in the phylogenetic analysis, indicating a potential language replacement from Afroasiatic to Niger-Congo in this population (Fig. 2). Prior studies suggest a complex history of the Fulani; analyses of Y chromosome variation suggest a shared ancestry with Nilo-Saharan and Afroasiatic populations [24], whereas mtDNA indicates a West African origin [26]. An analysis based on autosomal markers found traces of West Eurasian-related ancestry in this population [4], which suggests a North African or East African origin (as North and East Africans also have such ancestry likely related to expansions of farmers and herders from the Near East) and is consistent with the presence at moderate frequency of the −13,910T variant associated with lactose tolerance in European populations [15, 16].

Phylogenetic reconstruction of the relationship of African individuals under a model allowing for migration using TREEMIX [27] largely recapitulates the NJ phylogeny with the exception of the Fulani who cluster near neighboring Niger-Congo-speaking populations with whom they have admixed (Additional file 2: Figure S1). Interestingly, TREEMIX analysis indicates evidence for gene flow between the Hadza and the ancestors of the Ju|‘hoan and Khomani San, supporting genetic, linguistic, and archeological evidence that Khoesan-speaking populations may have originated in Eastern Africa [28, 29, 30].

afroasiatic-niger-congo-admixture
ADMIXTURE analysis of 92 African and 62 West Eurasian individuals. Each bar is an individual and colors represent the proportion of inferred ancestry from K ancestral populations. The bottom bar shows the language classification of each individual. With the increasing of K, the populations are largely grouped by their current language usage

About the Fulani, this is what the referenced study of Y‐chromosome variation among 15 Sudanese populations by Hassan et al. (2008), had to say:

  • Haplogroups A-M13 and B-M60 are present at high frequencies in Nilo-Saharan groups except Nubians, with low frequencies in Afro-Asiatic groups although notable frequencies of B-M60 were found in Hausa (15.6%) and Copts (15.2%).
  • Haplogroup E (four different haplotypes) accounts for the majority (34.4%) of the chromosome and is widespread in the Sudan. E-M78 represents 74.5% of haplogroup E, the highest frequencies observed in Masalit and Fur populations. E-M33 (5.2%) is largely confined to Fulani and Hausa, whereas E-M2 is restricted to Hausa. E-M215 was found to occur more in Nilo-Saharan rather than Afro-Asiatic speaking groups.
  • In contrast, haplogroups F-M89, I-M170, J-12f2, and JM172 were found to be more frequent in the Afro-Asiatic speaking groups. J-12f2 and J-M172 represents 94% and 6%, respectively, of haplogroup J with high frequencies among Nubians, Copts, and Arabs.
  • Haplogroup K-M9 is restricted to Hausa and Gaalien with low frequencies and is absent in Nilo-Saharan and Niger-Congo.
  • Haplogroup R-M173 appears to be the most frequent haplogroup in Fulani, and haplogroup R-P25 has the highest frequency in Hausa and Copts and is present at lower frequencies in north, east, and western Sudan.
  • Haplogroups A-M51, A-M23, D-M174, H-M52, L-M11, OM175, and P-M74 were completely absent from the populations analyzed.
fulfulde-fulani-language
Image modified from “Fulfulde Language Family Report” Author: Annette Harrison; Cartographer: Irene Tucker; SIL International 2003.

This is what David Reich will talk about in the seminar Insights into language expansions from ancient DNA:

In this talk, I will describe how the new science of genome-wide ancient DNA can provide insights into past spreads of language and culture. I will discuss five examples: (1) the spread of Indo-European languages to Europe and South Asia in association with Steppe pastoralist ancestry, (2) the spread of Austronesian languages to the open Pacific islands in association with Taiwanese aboriginal-associated ancestry, (3) the spread of Austroasiatic languages through southeast Asia in association with the characteristic ancestry type that is also represented in western Indonesia suggesting that these languages were once widespread there, (4) the spread of Afroasiastic languages through in East Africa as part of the Pastoral Neolithic farming expansion, and (5) the spread of Na-Dene languages in North America in association with Proto-Paleoeskimo ancestry. I will highlight the ways that ancient DNA can meaningfully contribute to our understanding of language expansions—increasing the plausibility of some scenarios while decreasing the plausibility of others—while emphasizing that with genetic data by itself we can never definitively determine what languages ancient people spoke.

EDIT (3 MAY 2019): Apparently, there was not much to take from the talk:

neolithic-pastoralist-africa
Pastoralist Neolithic in Africa, through a pale-green Sahelo-Sudanian steppe corridor. See full map.

This seminar (and maybe some new paper on the Neolithic expansion in Africa) could shed light on population movements that may be related to the spread of Afroasiatic dialects. Until now, it seems that Bantu peoples have been more interesting for linguistics and archaeology, and South and East Africans for anthropology.

Archaeology in Africa appears to be in its infancy, as is population genomics. From the latest publication by Carina Schlebusch, Population migration and adaptation during the African Holocene: A genetic perspective, a chapter from Modern Human Origins and Dispersal (2019):

The process behind the introduction and development of farming in Africa is still unclear. It is not known how many independent invention events there were in the continent and to which extent the various first instances of farming in northern Africa are linked. Based on the archeological record, it was proposed that at least three regions in Africa may have developed agriculture independently: the Sahara/Sahel (around 7 ka), the Ethiopian highlands (7-4 ka), and western Africa (5-3 ka). In addition to these developments, the Nile River Valley is thought to have adopted agriculture (around 7.2 ka), from the Neolithic Revolution in the Middle East (Chapter 12 – Jobling et al. 2014; Chapter 35, 37 – Mitchell and Lane 2013). From these diverse centers of origin, farmers or farming practices spread to the rest of Africa, with domesticate animals reaching the southern tip of Africa ~2 ka and crop farming ~1,8 ka (Mitchell 2002; Huffman 2007)

african-popularion-movements
Schematic representation of possible migration routes related to the expansion of herders and crop farmers during Holocene times. Arrow color indicate source populations; Brown-Eurasian, Green-western African, Blue-eastern African.

Similar to the case in Europe and the 1990s-2000s wrong haplogroup history based on the modern distribution of R1b, R1a, N, or I2, it is possible that neither of the most often mentioned haplogroups linked to the Afroasiatic expansion, E and J, were responsible for its early spread within Africa, despite their widespread distribution in certain modern Afroasiatic-speaking areas. The fact that such assessments include implausible glottochronological dates spanning up to 20,000 years for the parent language, combined with regional language continuities despite archaeological changes, makes them even more suspicious.

Similar to the case with Indo-Europeans and the “steppe ancestry” concept of the 2010s, it may be that the often-looked-for West Eurasian ancestry among Africans is the effect of recent migrations, unrelated to the Afroasiatic expansion. The results of this paper could be offering another sign of how this ancestry may have expanded only quite recently westwards from East Africa through the Sahel, after the Semitic expansion to the south:

1. From approximately 1000 BC, accompanying Nilo-Saharan peoples.

2. From approximately AD 1500, with the different population movements related to the nomadic Fulani:

sahel-nomadic-sedentary
Image from Sahel in West African History – Oxford Research Encyclopedia of African History.
  • Arguably, since the Fulani caste system wasn’t as elaborate in northern Nigeria, eastern Niger, and Cameroon, these specific groups would be a good example of the admixture with eastern populations, based on the (proportionally) huge amount of slaves they dealt with.
  • Similarly, it could be argued that the castes-based social stratification in most other territories (including Sudan) would have helped them keep a genetic make-up similar to their region of origin in terms of ancient lineages, hence similar to Chadic populations from west to east.

Reich’s assertion of the association of the language expansion with the spread of Pastoral Neolithic is still too vague, but – based on previous publications of ancient DNA in Africa and the Levant – I don’t have high hopes for a revolutionary paper in the near future. Without many samples and proper temporal transects, we are stuck with speculations based on modern distributions and scarce historical data.

fula-people-distribution
A distribution map of Fula people. Dark green: a major ethnic group; Medium: significant; Light: minor. Modified from image by Sarah Welch at Wikipedia.

About the potential genetic make-up of Cameroon before the arrival of the Neolithic, from the recent SAA 84th Annual Meeting (Abstracts in PDF):

Lipson, Mark (Harvard Medical School), Mary Prendergast (Harvard University), Isabelle Ribot (Université de Montréal), Carles Lalueza-Fox (Institute of Evolutionary Biology CSIC-UPF) and David Reich (Harvard Medical School)

[253] Ancient Human DNA from Shum Laka (Cameroon) in the Context of African Population History We generated genome-wide DNA data from four people buried at the site of Shum Laka in Cameroon between 8000–3000 years ago. One individual carried the deeply divergent Y chromosome haplogroup A00 found at low frequencies among some present-day Niger-Congo speakers, but the genome-wide ancestry profiles for all four individuals are very different from the majority of West Africans today and instead are more similar to West-Central African hunter-gatherers. Thus, despite the geographic proximity of Shum Laka to the hypothesized birthplace of Bantu languages and the temporal range of our samples bookending the initial Bantu expansion, these individuals are not representative of a Bantu source population. We present a phylogenetic model including Shum Laka that features three major radiations within Africa: one phase early in the history of modern humans, one close to the time of the migration giving rise to non-Africans, and one in the past several thousand years. Present-day West Africans and some East Africans, in addition to Central and Southern African hunter-gatherers, retain ancestry from the first phase, which is therefore still represented throughout the majority of human diversity in Africa today.

Related