Corded Ware—Uralic (IV): Hg R1a and N in Finno-Ugric and Samoyedic expansions

haplogroup-uralians

This is the fourth of four posts on the Corded Ware—Uralic identification:

Let me begin this final post on the Corded Ware—Uralic connection with an assertion that should be obvious to everyone involved in ethnolinguistic identification of prehistoric populations but, for one reason or another, is usually forgotten. In the words of David Reich, in Who We Are and How We Got Here (2018):

Human history is full of dead ends, and we should not expect the people who lived in any one place in the past to be the direct ancestors of those who live there today.

Haplogroup N

Another recurrent argument – apart from “Siberian ancestry” – for the location of the Uralic homeland is “haplogroup N”. This is as serious as saying “haplogroup R1” to refer to Indo-European migrations, but let’s explore this possibility anyway:

Ancient haplogroups

We have now a better idea of how many ancient migrations (previously hypothesized to be associated with westward Uralic migrations) look like in genetic terms. From Damgaard et al. (Science 2018):

These serial changes in the Baikal populations are reflected in Y-chromosome lineages (Fig. SA; figs. S24 to S27, and tables S13 and SI4). MAI carries the R haplogroup, whereas the majority of Baikal_EN males belong to N lineages, which were widely distributed across Northern Eurasia (29), and the Baikal_LNBA males all carry Q haplogroups, as do most of the Okunevo_EMBA as well as some present-day Central Asians and Siberians.

The only N1c1 sample comes from Ust’Ida Late Neolithic, 180km to the north of Lake Baikal, which – together with the Bronze Age sample from the Kola peninsula, and the medieval sample from Ust’Ida – gives a good idea of the overall expansion of N subclades and Siberian ancestry among the Circum-Arctic peoples of Eurasia, speakers of Palaeo-Siberian languages.

eurasian-n-subclades
Geographical location of ancient samples belonging to major clade N of the Y-chromosome.

Modern haplogroups

What we should expect from Uralic peoples expanding with haplogroup N – seeing how Yamna expands with R1b-L23, and Corded Ware expands with R1a-Z645 – is to find a common subclade spreading with Uralic populations. Let’s see if it works like that for any N-X subclade, in data from Ilumäe et al. (2016):

haplogroup_n1
Geographic-Distribution Map of hg N3 / N1c / N1a.

Within the Eurasian circum-Arctic spread zone, N3 and N2a reveal a well-structured spread pattern where individual sub-clades show very different distributions:

N1a1-M46 (or N-TAT), formed ca. 13900 BC, TMRCA 9800 BC

   N1a1a2-B187, formed ca. 9800 BC, TMRCA 1050 AD:

The sub-clade N3b-B187 is specific to southern Siberia and Mongolia, whereas N3a-L708 is spread widely in other regions of northern Eurasia.

     N1a1a1a-L708, formed ca. 6800 BC, TMRCA 5400 BC.

       N1a1a1a2-B211/Y9022, formed ca. 5400 BC, TMRCA 1900 BC:

The deepest clade within N3a is N3a1-B211, mostly present in the Volga-Uralic region and western Siberian Khanty and Mansi populations.

         N1a1a1a1a-L392/L1026), formed ca. 4400 BC, TMRCA 2800 BC:

The neighbor clade, N3a3’6-CTS6967, spreads from eastern Siberia to the eastern part of Fennoscandia and the Baltic States

haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders.

           N1a1a1a1a1a-CTS2929/VL29, formed ca. 2100 BC, TMRCA 1600 BC:

In Europe, the clade N3a3-VL29 encompasses over a third of the present-day male Estonians, Latvians, and Lithuanians but is also present among Saami, Karelians, and Finns (Table S2 and Figure 3). Among the Slavic-speaking Belarusians, Ukrainians, and Russians, about three-fourths of their hg N3 Y chromosomes belong to hg N3a3.

In the post on Finno-Permic expansions, I depicted what seems to me the most likely way of infiltration of N1c-L392 lineages with Akozino warrior-traders into the western Finno-Ugric populations, with an origin around the Barents sea.

This includes the potential spread of (a minority of) N1c-B211 subclades due to contacts with Anonino on both sides of the Urals, through a northern route of forest and forest-steppe regions (equivalent to the distribution of Cherkaskul compared to Andronovo), given the spread of certain subclades in Ugric populations.

NOTE. An alternative possibility is the association of certain B211 subclades with a southern route of expansion with Pre-Scythian and Scythian populations, under whose influence the Ananino culture emerged -which would imply a very quick infiltration of certain groups of haplogroup N everywhere among Finno-Ugrics on both sides of the Urals – , and also the expansion of some subclades with Turkic-speaking peoples, who apparently expanded with alliances of different peoples. Both (Scythian and Turkic) populations expanded from East Asia, where haplogroup N (including N1c) was present since the Neolithic. I find this a worse model of expansion for upper clades, but – given the YFull estimates and the presence of this haplogroup among Turkic peoples – it is a possibility for many subclades.

           N1a1a1a1a2-Z1936, formed ca. 2800 BC, TMRCA 2400 BC:

The only notable exception from the pattern are Russians from northern regions of European Russia, where, in turn, about two-thirds of the hg N3 Y chromosomes belong to the hg N3a4-Z1936—the second west Eurasian clade. Thus, according to the frequency distribution of this clade, these Northern Russians fit better among other non-Slavic populations from northeastern Europe. N3a4 tends to increase in frequency toward the northeastern European regions but is also somewhat unexpectedly a dominant hg N3 lineage among most Turcic-speaking Volga Tatars and South-Ural Bashkirs.

haplogroup_n3a4
Frequency-Distribution Maps of Individual Subclade N3a4 / N1a1a1a1a2-Z1936, probably with the Samic (first) and Fennic (later) expansions into Paleo-Lakelandic and Palaeo-Laplandic territories.

The expansion of N1a-Z1936 in Fennoscandia is most likely associated with the expansion of Saami into asbestos ware-related territory (like the Lovozero culture) during the Late Iron Age – and mixture with its population – , and with the later Fennic expansion to the east and north, replacing their language.

           N1a1a1a1a4-M2019 (previously N3a2), formed ca. 4400 BC, TMRCA 1700 BC:

Sub-hg N3a2-M2118 is one of the two main bifurcating branches in the nested cladistic structure of N3a2’6-M2110. It is predominantly found in populations inhabiting present-day Yakutia (Republic of Sakha) in central Siberia and at lower frequencies in the Khanty and Mansi populations, which exhibit a distinct Y-STR pattern (Table S7) potentially intrinsic to an additional clade inside the sub-hg N3a2

The second widespread sub-clade of hg N is N2a. (…):

   N1a2b-P43 (B523/FGC10846/Y3184), formed ca. 6800 BC, TMRCA ca. 2700 BC:

The absolute majority of N2a individuals belong to the second sub-clade, N2a1-B523, which diversified about 4.7 kya (95% CI = 4.0–5.5 kya). Its distribution covers the western and southern parts of Siberia, the Taimyr Peninsula, and the Volga-Uralic region with frequencies ranging from from 10% to 30% and does not extend to eastern Siberia (…)

haplogroup_n2
Geographic-Distribution Map of hg N2a1 / N1a2b-P43

The “European” branch suggested earlier from Y-STR patterns turned out to consist of two clades

     N1a2b2a-Y3185/FGC10847, formed ca. 2200 BC, TMRCA 800 BC:

N2a1-L1419, spread mainly in the northern part of that region.

     N1a2b2b1-B528/Y24382, formed ca. 900 BC, TMRCA ca. 900 BC:

N2a1-B528, spread in the southern Volga-Uralic region.

Haplogroup R1a

We also have a good idea of the distribution of haplogroup R1a-Z645 in ancient samples. Its subclades were associated with the Corded Ware expansion, and some of them fit quite well the early expansion of Finno-Permic, Ugric, and Samoyedic peoples to the east.

r1a-z282-z280-z2125-distribution
Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups.. Notice the potential Finno-Ugric-associated distribution of Z282 (especially R1a-M558, a Z280 subclade), the expansion of R1a-Z2123 subclades with Central Asian forest-steppe groups.

This is how the modern distribution of R1a among Uralians looks like, from the latest report in Tambets et al. (2018):

  • Among Fennic populations, Estonians and Karelians (ca. 1.1 million) have not suffered the greatest bottleneck of Finns (ca. 6-7 million), and show thus a greater proportion of R1a-Z280 than N1c subclades, which points to the original situation of Fennic peoples before their expansion. To trust Finnish Y-DNA to derive conclusions about the Uralic populations is as useful as relying on the Basque Y-DNA for the language spread by R1b-P312
  • Among Volga-Finnic populations, Mordovians (the closest to the original Uralic cluster, see above) show a majority of R1a lineages (27%).
  • Hungarians (ca. 13-15 million) represent the majority of Ugric (and Finno-Ugric) peoples. They are mainly R1a-Z280, also R1a-Z2123, have little N1c, and lack Siberian ancestry, and represent thus the most likely original situation of Ugric peoples in 4th century AD (read more on Avars and Hungarians).
  • Among Samoyedic peoples, the Selkup, the southernmost ones and latest to expand – that is, those not heavily admixed with Siberian populations – , also have a majority of R1a-Z2123 lineages (see also here for the original Samoyedic haplogroups to the south).

To understand the relevance of Hungarians for Ugric peoples, as well as Estonians, Karelians, and Mordovians (and northern Russians, Finno-Ugric peoples recently Russified) for Finno-Permic peoples, as opposed to the Circum-Arctic and East Siberian populations, one has to put demographics in perspective. Even a modern map can show the relevance of certain territories in the past:

population-density
Population density (people per km2) map of the world in 1994. From Wikipedia.

Summary of ancestry + haplogroups

Fennic and Samic populations seem to be clearly influenced by Palaeo-Laplandic peoples, whereas Volga-Finnic and especially Permic populations may have received gene flow from both, but essentially Palaeo-Siberian influence from the north and east.

The fact that modern Mansis and Khantys offer the highest variation in N1a subclades, and some of the highest “Siberian ancestry” among non-Nganasans, should have raised a red flag long ago. The fact that Hungarians – supposedly stemming from a source population similar to Mansis – do not offer the same amount of N subclades or Siberian ancestry (not even close), and offer instead more R1a, in common with Estonians (among Finno-Samic peoples) and Mordvins (among Volga-Finnic peoples) should have raised a still bigger red flag. The fact that Nganasans – the model for Siberian ancestry – show completely different N1a2b-P43 lineages should have been a huge genetic red line (on top of the anthropological one) to regard them as the Uralian-type population.

We know now that ethnolinguistic groups have usually expanded with massive (usually male-biased) migrations, and that neighbouring locals often ‘resurge’ later without changing the language. That is seen in Europe after the spread of Bell Beakers, with the increase of previous ancestry and lineages in Scandinavia during the formation of the Nordic ethnolinguistic community; in Central-West Europe, with the resurgence of Neolithic ancestry (and lineages) during the Bronze Age over steppe ancestry; and in Central-East Europe (with Unetice or East European Bronze Age groups like Mierzanowice, Trzciniec, or Lusatian) showing an increase in steppe ancestry (and resurge of R1a subclades); none of them represented a radical ethnolinguistic change.

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).

It is not hard to model the stepped arrival, infiltration, and/or resurge of N subclades and “Siberian ancestries”, as well as their gradual expansion in certain regions, associated with certain migrations first – such as the expansions to the Circum-Arctic region, and later the Scythian- and Turkic-related movements – , as well as limited regional developments, like the known bottleneck in Finns, or the clear late expansion of Ugric and Samoyedic languages to the north among nomadic Palaeo-Siberians due to traditions of exogamy and multilingualism. This fits quite well with the different arrival of N (N1c and xN1c) lineages to the different Uralic-speaking groups, and to the stepped appearance of “Siberian ancestry” in the different regions.

The aternative

It is evident that a lot of people were too attached to the idea of Palaeolithic R1b lineages ‘native’ to western Europe speaking Basque languages; of R1a lineages speaking Indo-European and spreading with Yamna; and N lineages ‘native’ to north-eastern Europe and speaking Uralic, and this is causing widespread weeping and gnashing of teeth (instead of the joy of discovering where one’s true patrilineal ancestors come from, and what language they spoke in each given period, which is the supposed objective of genetic genealogy…)

Since an Indo-Germanic branch (as revived now by some in the Copenhaguen group to fit Kristiansen’s theory of the 1980s with recent genetic data) does not make any sense in linguistics, the finding of R1a in Yamna would not have led where some think it would have, because North-West Indo-European would still be the main Late PIE branch in Europe. Don’t take my word for it; take James P. Mallory’s (2013).

mallory-adams-tree
The levels of Indo-European reconstruction, from Mallory & Adams (2006).

If an (unlikely) Indo-Slavonic group were posited, though, such a group would still be bound (with Indo-Iranian) to the steppes with East Yamna/Poltavka (admixing with Abashevo migrants, but retaining its language), developing Sintashta/Potapovka → Srubna/Andronovo, and R1a lineages would have equally undergone the known bottlenecks of the steppes where they replaced R1b-Z2103 – which this eastern group shares with Balkan languages, a haplogroup that links therefore together the Graeco-Aryan group.

As far as I know – and there might be many other similar pet theories out there – there have been proposals of “modern Balto-Slavic-like” populations (in an obvious circular reasoning based on modern populations) in some Scythian clusters of the Iron Age.

NOTE. I will not enter into “Balto-Slavic-like R1a” of the Late Bronze Age or earlier because no one can seriously believe at this point of development of Population Genetics that autosomal similarity predating 1,500+ years the appearance of Slavs equates to their (ethnolinguistic) ancestral population, without a clear intermediate cultural and genetic trail – something we lack today in the Slavic case even for the late Roman period…

finno-saamic-palaeo-germanic-substratum
The Finnic and Saamic separation looks shallower than it actually is. Invisible convergence can be ‘triangulated’ with the help of Germanic layers of mutual loanwords (Häkkinen 2012).

We also know of R1a-Z280 lineages in Srubna, probably expanding to the west. With that in mind, and knowing that Palaeo-Germanic was in close contact with Finno-Samic while both were already separated but still in contact, and that Palaeo-Germanic was also in contact and closely related to a ‘Temematic’ distinct from Balto-Slavic (and also that early Proto-Baltic and Proto-Slavic from the Roman Iron Age and later were in contact with western Uralic) this will be the linguistic map of the Iron Age if R1a is considered to expand Indo-European from some kind of “patron-client” relationship with west Yamna:

palaeo-germanic-italo-celtic
Eastern European language map during the Late Bronze Age / Iron Age, if R1a spread Indo-European languages and Eastern Yamna spoke Indo-Slavonic. Palaeo-Germanic (i.e. Pre- to Proto-Germanic) needs to be in contact with both the Samic Lovozero population and the Fennic west Circum-Arctic one. Italic and Celtic in contact with Pre-Germanic. Germanic in contact with Temematic. Balto-Slavic in contact with Iranian, and near Fennic to allow for later loanwords. For Germanic and Temematic, see Kortlandt (2018).

You might think I have some personal or political reason against this kind of proposals. I haven’t. We have been proposing Indo-European to be the language of the European Union for more than 10 years, so to support R1b-Italo-Celtic in the whole Western Europe, R1a-Germanic in Central and Eastern Europe, and R1a-Indo-Slavonic in the steppes (as the Danish group seems to be doing) has nothing inherently bad (or good) for me. If anything, it gives more reason to support the revival of North-West Indo-European in Europe.

My problem with this proposal is that it is obviously beholden to the notion of the uninterrupted cultural, historic and ethnic continuity in certain territories. This bias is common in historiography (von Falkenhausen 1993), but it extends even more easily into the lesser known prehistory of any territory, and now more than ever some people feel the need to corrupt (pre)history based on their own haplogroups (or the majority haplogroups of their modern countries). However, more than on philosophical grounds, my rejection is based on facts: this picture is not what the combination of linguistic, archaeological, and genetic data shows. Period.

Nevertheless, if Yamna + Corded Ware represented the “big and early expansion” of Germanic and Italo-Celtic peoples proper of the dream Nazi’s Lebensraum and Fascist’s spazio vitale proposals; Uralians were Siberian hunter-gatherers that controlled the whole eastern and northern Russia, and miraculously managed to push (ethnolinguistically) Neolithic agropastoralists to the west during and after the Iron Age, with gradual (and often minimal) genetic impact; and Balto-Slavic peoples were represented by horse riders from Pokrovka/Srubna, hiding then somewhere around the forest-steppe until after the Scythian expansion, and then spreading their language (without much genetic impact) during the early Middle Ages…so be it.

Related

Corded Ware—Uralic (III): “Siberian ancestry” and Ugric-Samoyedic expansions

siberian-ancestry-tambets

This is the third of four posts on the Corded Ware—Uralic identification. See

An Eastern Uralic group?

Even though proposals of an Eastern Uralic (or Ugro-Samoyedic) group are in the minority – and those who support it tend to search for an origin of Uralic in Central Asia – , there is nothing wrong in supporting this from the point of view of a western homeland, because the eastward migration of both Proto-Ugric and Pre-Samoyedic peoples may have been coupled with each other at an early stage. It’s like Indo-Slavonic: it just doesn’t fit the linguistic data as well as the alternative, i.e. the expansion of Samoyedic first, different from a Finno-Ugric trunk. But, in case you are wondering about this possibility, here is Häkkinen’s (2012) phonological argument:

ugro-samoyedic-uralic

The case of Samoyedic is quite similar to that of Hungarian, although the earliest Palaeo-Siberian contact languages have been lost. There were contacts at least with Tocharian (Kallio 2004), Yukaghir (Rédei 1999) and Turkic (Janhunen 1998). Samoyedic also:

a) has moved far from the related languages and has been exposed to strong foreign influence

b) shares a small number of common words with other branches (from Sammallahti 1988: only 123 ‘Uralic’ words, versus 390 ‘Uralic’ + ‘Finno-Ugric’ words found in other branches than Samoyedic = 31,5 %)

c) derives phonologically from the East Uralic dialect.

The phonological level is taxonomically more reliable, since it lacks the distortion caused by invisible convergence and false divergence at the lexical level. Thus we can conclude that the traditional taxonomic model, according to which Samoyedic was the first branch to split off from the Proto-Uralic unity, is just as incorrect as the view that Hungarian was the first branch to split off.

Seima-Turbino

Late Uralic can be traced back to metallurgical cultures thanks to terms like PU *wäśka ‘copper/bronze’ (borrowed from Proto-Samoyedic *wesä into Tocharian); PU *äsa and *olna/*olni, ‘lead’ or ‘tin’, found in *äsa-wäśka ‘tin-bronze’; and e.g. *weŋći ‘knife’, borrowed into Indo-Iranian (through the stage of vocalization of nasals), appearing later as Proto-Indo-Aryan *wāćī ‘knife, awl, axe’.

It is known that the southern regions of the Abashevo culture developed Proto-Indo-Iranian-speaking Sintashta-Petrovka and Pokrovka (Early Srubna). To the north, however, Abashevo kept its Uralic nature, with continuous contacts allowing for the spread of lexicon – mainly into Finno-Ugric – , and phonetic influence – mainly Uralisms into Proto-Indo-Iranian phonology (read more here).

The northern part of Abashevo (just like the south) was mainly a metallurgical society, with Abashevo metal prospectors found also side by side with Sintashta pioneers in the Zeravshan Valley, near BMAC, in search of metal ores. About the Seima-Turbino phenomenon, from Parpola (2013):

From the Urals to the east, the chain of cultures associated with this network consisted principally of the following: the Abashevo culture (extending from the Upper Don to the Mid- and South Trans-Urals, including the important cemeteries of Sejma and Turbino), the Sintashta culture (in the southeast Urals), the Petrovka culture (in the Tobol-Ishim steppe), the Taskovo-Loginovo cultures (on the Mid- and Lower Tobol and the Mid-Irtysh), the Samus’ culture (on the Upper Ob, with the important cemetery of Rostovka), the Krotovo culture (from the forest steppe of the Mid-Irtysh to the Baraba steppe on the Upper Ob, with the important cemetery of Sopka 2), the Elunino culture (on the Upper Ob just west of the Altai mountains) and the Okunevo culture (on the Mid-Yenissei, in the Minusinsk plain, Khakassia and northern Tuva). The Okunevo culture belongs wholly to the Early Bronze Age (c. 2250–1900 BCE), but most of the other cultures apparently to its latter part, being currently dated to the pre-Andronovo horizon of c. 2100–1800 BCE (cf. Parzinger 2006: 244–312 and 336; Koryakova & Epimakhov 2007: 104–105).

post-eneolithic-steppe-asia
Schematic map of the Middle Bronze Age cultures (steppe and foreststeppe
zone)

The majority of the Sejma-Turbino objects are of the better quality tin-bronze, and while tin is absent in the Urals, the Altai and Sayan mountains are an important source of both copper and tin. Tin is also available in southern Central Asia. Chernykh & Kuz’minykh have accordingly suggested an eastern origin for the Sejma-Turbino network, backing this hypothesis also by the depiction on the Sejma-Turbino knives of mountain sheep and horses characteristic of that area. However, Christian Carpelan has emphasized that the local Afanas’evo and Okunevo metallurgy of the Sayan-Altai area was initially rather primitive, and could not possibly have achieved the advanced and difficult technology of casting socketed spearheads as one piece around a blank. Carpelan points out that the first spearheads of this type appear in the Middle Bronze Age Caucasia c. 2000 BCE, diffusing early on to the Mid-Volga-Kama-southern Urals area, where “it was the experienced Abashevo craftsmen who were able to take up the new techniques and develop and distribute new types of spearheads” (Carpelan & Parpola 2001: 106, cf. 99–106, 110). The animal argument is countered by reference to a dagger from Sejma on the Oka river depicting an elk’s head, with earlier north European prototypes (Carpelan & Parpola 2001: 106–109). Also the metal analysis speaks for the Abashevo origin of the Sejma-Turbino network. Out of 353 artefacts analyzed, 47% were of tin-bronze, 36% of arsenical bronze, and 8.5% of pure copper. Both the arsenical bronze and pure copper are very clearly associated with the Abashevo metallurgy.

seima-turbino-phenomenon-parpola
Find spots of artefacts distributed by the Sejma-Turbino intercultural trader network, and the areas of the most important participating cultures: Abashevo, Sintashta, Petrovka. Based on Chernykh 2007: 77.

The Abashevo metal production was based on the Volga-Kama-Belaya area sandstone ores of pure copper and on the more easterly Urals deposits of arsenical copper (Figure 9). The Abashevo people, expanding from the Don and Mid-Volga to the Urals, first reached the westerly sandstone deposits of pure copper in the Volga and Kama basins, and started developing their metallurgy in this area, before moving on to the eastern side of the Urals to produce harder weapons and tools of arsenical copper. Eventually they moved even further south, to the area richest in copper in the whole Urals region, founding there the very strong and innovative Sintashta culture.

Regarding the most likely expansion of Eastern Uralic peoples:

Nataliya L’vovna Chlenova (1929–2009; cf. Korenyako & Ku’zminykh 2011) published in 1981 a detailed study of the Cherkaskul’ pottery. In her carefully prepared maps of 1981 and 1984 (Figure 10), she plotted Cherkaskul’ monuments not only in Bashkiria and the Trans-Urals, but also in thick concentrations on the Upper Irtysh, Upper Ob and Upper Yenissei, close to the Altai and Sayan mountains, precisely where the best experts suppose the homeland of Proto-Samoyed to be.

cherkaskul-andronovo
Distribution of Srubnaya (Timber Grave, early and late), Andronovo (Alakul’ and Fëdorovo variants) and Cherkaskul’ monuments. After Parpola 1994: 146, fig. 8.15, based on the work of N. L. Chlenova (1984: map facing page 100).

Ugric

The Cherkaskul’ culture was transformed into the genetically related Mezhovka culture (c. 1500–1000 BCE), which occupied approximately the same area from the Mid-Kama and Belaya rivers to the Tobol river in western Siberia (cf. Parzinger 2006: 444–448; Koryakova & Epimakhov 2007: 170–175). The Mezhovka culture was in close contact with the neighbouring and probably Proto-Iranian speaking Alekseevka alias Sargary culture (c. 1500–900 BCE) of northern Kazakhstan (Figure 4 no. 8) that had a Fëdorovo and Cherkaskul’ substratum and a roller pottery superstratum (cf. Parzinger 2006: 443–448; Koryakova & Epimakhov 2007: 161–170). Both the Cherkaskul’ and the Mezhovka cultures are thought to have been Proto-Ugric linguistically, on the basis of the agreement of their area with that of Mansi and Khanty speakers, who moreover in their Fëdorovo-like ornamentation have preserved evidence of continuity in material culture (cf. Chlenova 1984; Koryakova & Epimakhov 2007: 159, 175).

mezhovska-sargary-irmen
Cultures of the Final Bronze Age of the Urals and western Siberia (steppe
and forest-steppe zone).

The Mezhovka culture was succeeded by the genetically related Gamayun culture (c. 1000–700 BCE) (cf. Parzinger 2006: 446; 542–545).

From the Gamayun culture descend Trans-Urals cultures in close contact with Finno-Permic populations of the Cis-Ural region:

  • [Proto-Mansi] Itkul’ culture (c. 700–200 BCE) distributed along the eastern slope of the Ural Mountains (cf. Parzinger 2006: 552–556). Known from its walled forts, it constituted the principal Trans-Uralian centre of metallurgy in the Iron Age, and was in contact with both the Anan’ino and Akhmylovo cultures (the metallurgical centres of the Mid-Volga and Kama-Belaya region) and the neighbouring Gorokhovo culture.
    • [Proto-Hungarian] via the Vorob’evo Group (c. 700–550 BCE) (cf. Parzinger 2006: 546–549), to the Gorokhovo culture (c. 550–400 BCE) of the Trans-Uralian forest steppe (cf. Parzinger 2006: 549–552). For various reasons the local Gorokhovo people started mobile pastoral herding and became part of the multicomponent pastoralist Sargat culture (c. 500 BCE to 300 CE), which in a broader sense comprized all cultural groups between the Tobol and Irtysh rivers, succeeding here the Sargary culture. The Sargat intercommunity was dominated by steppe nomads belonging to the Iranian-speaking Saka confederation, who in the summer migrated northwards to the forest steppe
  • [Proto-Khanty] Late Bronze Age and Early Iron Age cultures related to the Gamayunskoe and Itkul’ cultures that extended up to the Ob: the Nosilovo, Baitovo, Late Irmen’, and Krasnoozero cultures (c. 900–500 BCE). Some were in contact with the Akhmylovo on the Mid-Volga.
sargat-gorokhovo-bolscherechye
Cultural groups of the Iron Age in the forest-steppe zone of western
Siberia. (

Samoyedic

Parpola (2012) connects the expansion of Samoyedic with the Cherkaskul variant of Andronovo. As we know, Andronovo was genetically diverse, which speaks in favour of different groups developing similar material cultures in Central Asia.

Juha Janhunen, author of the etymological dictionary of the Samoyed languages (1977), places the homeland of Proto-Samoyedic in the Minusinsk basin on the Upper Yenissei (cf. Janhunen 2009: 72). Mainly on the basis of Bulghar Turkic loanwords, Janhunen (2007: 224; 2009: 63) dates Proto-Samoyedic to the last centuries BCE. Janhunen thinks that the language of the Tagar culture (c. 800–100 BCE) ought to have been Proto-Samoyedic (cf. Janhunen 1983: 117– 118; 2009: 72; Parzinger 2001: 80 and 2006: 619–631 dates the Tagar culture c. 1000–200 BCE; Svyatko et al. 2009: 256, based on human bone samples, c. 900 BCE to 50 CE). The Tagar culture largely continues the traditions of the Karasuk culture (c. 1400–900 BCE), (…)

chicha-irmen-tagar-baraba-forest-siberian
Map showing the location of Chicha-1.

For the most recent expansions of Samoyedic languages to the north, into Palaeo-Siberian populations, read more about the traditional multilingualism of Siberian populations.

Genetics

Siberian ancestry

The use of a map of “Siberian ancestry” peaking in the arctic to show a supposedly late Uralic population movement (starting in the Iron Age!) seems to be the latest trend in population genomics:

siberian-ancestry-map
Frequency map of the so-called ‘Siberian’ component. From Tambets et al. (2018) (see below for ADMIXTURE in specific populations).

I guess that would make this map of Neolithic farmer ancestry represent an expansion of Indo-European from the south, because Anatolia, Greece, Italy, southern France, and Iberia – where this ancestry peaks in modern populations – are among the oldest territories where Indo-European languages were recorded:

reich-farmer-ancestry
Modern genome-wide data shows that the primary gradient of farmer ancestry in Europe does not flow southeast-to-northwest but instead in an almost perpendicular direction, a result of a major migration of pastoralists from the east that displaced much of the ancestry of the first farmers.

Probably not the right interpretation of this kind of simplistic data about modern populations, though…

The most striking thing about the “Siberian ancestry” white whale is that nobody really knows what it is; just like we did not know what “Yamnaya ancestry” was, until the most recent data is making the picture clearer. Its nature is changing with each new paper, and it can be summed up by “some ancestry we want to find that is common to Uralic-speaking peoples, and should not be CWC-related”. Tambets et al. (2018) explain quite well how they “found it”:

Overall, and specifically at lower values of K, the genetic makeup of Uralic speakers resembles that of their geographic neighbours. The Saami and (a subset of) the Mansi serve as exceptions to that pattern being more similar to geographically more distant populations (Fig. 3a, Additional file 3: S3). However, starting from K = 9, ADMIXTURE identifies a genetic component (k9, magenta in Fig. 3a, Additional file 3: S3), which is predominantly, although not exclusively, found in Uralic speakers. This component is also well visible on K = 10, which has the best cross-validation index among all tests (Additional file 3: S3B). The spatial distribution of this component (Fig. 3b) shows a frequency peak among Ob-Ugric and Samoyed speakers as well as among neighbouring Kets (Fig. 3a). The proportion of k9 decreases rapidly from West Siberia towards east, south and west, constituting on average 40% of the genetic ancestry of FU speakers in Volga-Ural region (VUR) and 20% in their Turkic-speaking neighbours (Bashkirs, Tatars, Chuvashes; Fig. 3a).

siberian-ancestry-modern
Population structure of Uralic-speaking populations inferred from ADMIXTURE analysis on autosomal SNPs in Eurasian context. Individual ancestry estimates for populations of interest for selected number of assumed ancestral populations (K3, K6, K9, K11). Ancestry components discussed in a main text (k2, k3, k5, k6, k9, k11) are indicated and have the same colours throughout. The names of the Uralic-speaking populations are indicated with blue (Finno-Ugric) or orange (Samoyedic). Image from Tambets et al. (2018).

However, this ‘something’ that some people occasionally find in some Uralic populations is also common to other modern and ancient groups, and not so common in some other Uralic peoples. Simply put:

siberian-ancestry-modern-populations
Image modified from Lamnidis et al. (2018). Red line representing maximum “Siberian admixture” in Eastern European hunter-gatherers. In blue, Uralic-speaking groups. “Plot of ADMIXTURE (K=3) results containing West Eurasian populations and the Nganasan. Ancient individuals from this study are represented by thicker bars.”

I already said this in the recent publication of Siberian samples, where a renamed and radiocarbon dated Finnish_IA clearly shows that Late Iron Age Saami (ca. 400 AD) had little “Siberian ancestry”, if any at all, representing the most likely Fennic (and Samic) ancestral components before their expansion into central and northern Finland, where they admixed with circum-polar peoples of asbestos ware cultures.

I will say that again and again, any time they report the so-called “Siberian ancestry” in Uralic samples, no matter how it is defined each time: it does not seem to be that special something people are looking for, but rather (at least in a great part) a quite old ancestral component forming an evident cline with EHG, whose best proximate source are Baikal_EN (and/or Devil’s Gate) at this moment, and thus also East European hunter-gatherers for Western Uralic peoples:

dzudzuana-baikal-en-admixture
Image modified from Lazaridis et al. (2018). In red: samples with Baikal_EN ancestry in speculative estimates. In pink: samples with Baikal_EN ancestry in conservative estimates (probably marking a recent arrival of Baikal_En ancestry, see here). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (Left) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown. (Right) ‘Speculative’ estimates. The highest number of sources (≤5) with admixture estimates within [0,1] are shown for each population. Some of the admixture proportions are not significantly different from 0 (Supplementary Information section 4).

So either Samara_HG, Karelia_HG, and many other groups from eastern Europe all spoke Uralic according to this ADMIXTURE graphic (and the formation of steppe ancestry in the Volga-Ural region brought the Proto-Indo-European language to the steppes through the CHG/ANE expansion), or a great part of this “Siberian ancestry” found in modern Uralic-speaking populations is not what some people would like to think it is…

Modern populations

PCA clines can be looked for to represent expansions of ancient populations. Most recently, Flegontov et al. (2018) are attempting to do this with Asian populations:

For some Turkic groups in the Urals and the Altai regions and in the Volga basin, a different admixture model fits the data: the same West Eurasian source + Uralic- or Yeniseian-speaking Siberians. Thus, we have revealed an admixture cline between Scythians and the Iranian farmer genetic cluster, and two further clines connecting the former cline to distinct ancestry sources in Siberia. Interestingly, few Wusun-period individuals harbor substantial Uralic/Yeniseian-related Siberian ancestry, in contrast to preceding Scythians and later Turkic groups characterized by the Tungusic/Mongolic-related ancestry. It remains to be elucidated whether this genetic influx reflects contacts with the Xiongnu confederacy. We are currently assembling a collection of samples across the Eurasian steppe for a detailed genetic investigation of the Hunnic confederacies.

jeong-population-clines
Three distinct East/West Eurasian clines across the continent with some interesting linguistic correlates, as earlier reported by Jeong et al. (2018). Alexander M. Kim.

There are potential errors with this approach:

The main one is practical – does a modern cline represent an ancestral language? The answer is: sometimes. It depends on the anthropological context that we have, and especially on the precision of the PCA:

clines-himalayan
Genetic structure of the Himalayan region populations from analyses using unlinked SNPs. (A) PCA of the Himalayan and HGDP-CEPH populations. Each dot represents a sample, coded by region as indicated. The Himalayan region samples lie between the HGDP-CEPH East Asian and South Asian samples on the right-hand side of the plot. From Arciero et al. (2018).

The ‘Europe’, ‘Middle East’, etc. clines of the above PCA do not represent one language, but many. For starters, the PCA includes too many (and modern) populations, its precision is useless for ethnolinguistic groups. Which is the right level? Again, it depends.

The other error is one of detail of the clines drawn (which, in turn, depends on the precision of the PCA). For example, we can draw two paralell lines (or even one line, as in Flegontov et al. above) in one PCA graphic, but we still don’t have the direction of expansion. How do we know if this supposed “Uralic-speaking cline” goes from one region to the other? For that level of detail, we should examine closely modern Uralic-speaking peoples and Circum-Arctic populations:

uralic-cline
Modified from Tambets et al. (2018). Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations

The real ancient Uralic cluster (drawn above in blue) is thus probably from a North-East European source (probably formed by Battle Axe / Fatyanovo-Balanovo / Abashevo) to the east into Siberian populations, and to the north into Laplandic populations (see below also on Mezhovska ancestry for the drawn ‘European cline’, which some may a priori wrongly assume to be quite late).

The fact that the three formed clines point to an admixture of CWC-related populations from North-Eastern Europe, and that variation is greater at the Palaeo-Laplandic and Palaeo-Siberian extremities compared to the CWC-related one, also supports this as the correct interpretation.

However, judging by the two main clines formed, one could be alternatively inclined to interpret that Palaeo-Laplandic and Palaeo-Siberian populations formed a huge ancestral “Uralic” ghost cluster in Siberia (spanning from the Palaeo-Laplandic to the Palaeo-Siberian one), and from there expanded Finno-Samic on one hand, and “Volga-Ugro-Samoyed” on the other. That poses different problems: an obvious linguistic and archaeological one – which I assume a lot of people do not really care about – , and a not-so-obvious genetic one (see below for ancient samples and for the expansion of haplogroup N).

To understand the simplest solution better, one can just have a look at the PCA from Bell Beaker samples in Olalde et al. (2018), which (as Reich has already explained many times) expanded directly from Yamna R1b-L23 lineages:

olalde_pca_clines
Image modified from Olalde et al. (2018). PCA of 999 Eurasian individuals. Marked is the Espersted Outlier with the approximate position of Yamna Hungary, probably the source of its admixture. Different Bell Beaker clines have been drawn, to represent approximate source of expansions from Central European sources into the different regions.

Unlike this PCA with ancient samples, where Bell Beaker clines could be a rough approximation to the real sources for each population, and where a cluster spanning all three depicted Early Bronze Age clusters could give a rough proximate source of European Bell Beakers in Hungary (and where one can even distinguish the Y-DNA bottlenecks in the L23 trunk created by each cline) the PCA of modern Uralic populations is probably not suitable for a good estimate of the ancient situation, which may be found shifted up or down of the drawn “Uralic” cluster along East European groups.

After all, we already know that the Siberian cline shows probably as much an ancient admixture event – from the original Uralic expansion to the east with Corded Ware ancestry – as another more recent one – a westward migration of Siberian ancestry (or even more than one). While we know with more or less exactitude what happened with the Palaeo-Laplandic admixture by expanding Proto-Finno-Samic populations (see here), the Proto-Ugric and Pre-Samoyedic populations formed probably more than one cline during the different ancient migrations through central Asia.

Ancient populations

Apparently, the Corded Ware expansion to the east was not marked by a huge change in ancestry. While the final version of Narasimhan et al. (2018) may show a little more detail about other forest-steppe Seima-Turbino/Andronovo-related migrations (and thus also Eastern Uralic peoples), we have already had enough information for quite some time to get a good idea.

mezhovska-pca
Principal component analysis. PCA of ancient individuals (according colours see legend) projected on modern West Eurasians (grey). Iron Age Scythians are shown in black; CHG, Caucasus hunter-gatherer; LNBA, late Neolithic/Bronze Age; MN, middle Neolithic; EHG, eastern European huntergatherer; LBK_EN, early Neolithic Linearbandkeramik; HG, hunter-gatherer; EBA, early Bronze Age; IA, Iron Age; LBA, late Bronze Age; WHG, western hunter-gatherer.dataset (grey). Iron Age Scythians are shown in black; CHG, Caucasus hunter-gatherer; LNBA, late Neolithic/Bronze Age; MN, middle Neolithic; EHG, eastern European hunter-gatherer; LBK_EN, early Neolithic Linearbandkeramik; HG, hunter-gatherer; EBA, early Bronze Age; IA, Iron Age; LBA, late Bronze Age; WHG, western hunter-gatherer.

Mezhovska‘s position is similar to the later Pre-Scythian and Scythian populations. There are some interesting details: apart from haplogroup R1a-Z280 (CTS1211+), there is one R1b-M269 (PF6494+), probably Z2103, and an outlier (out of three) in a similar position to the recently described central/southern Scythian clusters.

NOTE. The finding of R1b-M269 in the forest-steppe is probably either 1) from an Afanasevo-Okunevo origin, or 2) from an admixture with neighbouring Andronovo-related populations, such as Sargary. A third, maybe less likely option is that this haplogroup admixed with Abashevo directly (as it happened in Sintashta, Potapovka, or Pokrovka) and formed part of early Uralic migrations. In any case, since Mezhovska is a Bronze Age society from the Urals region, its association with R1b-Z2103 – like the association of R1b-Z2103 in Scythian clusters – cannot be attributed to “Thracian peoples”, a link which is (as I already said) too simplistic.

The drawn “European cline” of Hungarians (see above), leading from ‘west-like’ Mansi to Hungarian populations – and hosting also Finnic and Estonian samples – , cannot therefore be attributed simply to late “Slavic/Balkan-like” admixture.

Karasuk – located further to the east – is basically also Corded Ware peoples showing clearly a recent admixture with local ANE / Baikal_EN-like populations. In terms of haplogroups it shows haplogroup Q, R1a-Z2124, and R1a-Z2123, later found among early Hungarians, and present also in ancient Samoyedic populations now acculturated.

The most interesting aspect of both Mezhovska and Karasuk is that they seem to diverge from a point close to Ukraine_Eneolithic, which is the supposed ancestral source of Corded Ware peoples (read more about the formation of “steppe ancestry”). This means that Eastern Uralians derive from a source closer to Middle Dnieper/Abashevo populations, rather than Battle Axe (shifted to Latvian Neolithic), which is more likely the source prevalent in Finno-Permic peoples.

Their initial admixture with (Palaeo-)Siberian populations is thus seen already starting by this time in Mezhovska and especially in Karasuk, but this process (compared to modern populations) is incomplete:

f4-test-karasuk-mezhovska
Visualization of f-statistics results. f4(Test, LBK; Han, Mbuti) values are plotted on x axis and f4(Test, LBK; EHG, Mbuti) values on y axis, positive deviations from zero show deviations from a clade between Test and LBK. A red dashed line is drawn between Yamnaya from Samara and Ami. Iron Age populations that can be modelled as mixtures of Yamnaya and East Eurasians (like the Ami) are arrayed around this line and appear to be distinct from the main North/South European cline (blue) on the left of the x axis.
karasuk-mezhovska-admixture
ADMIXTURE results for ancient populations. Red arrows point to the Iron Age Scythian individuals studied. LBK_EN: Early Neolithic Linearbandkeramik; EHG: Eastern European hunter-gatherer; Motala_HG: hunter-gatherer from Motala (Sweden); WHG: western hunter-gatherer; CHG: Caucasus hunter-gatherer; IA: Iron Age; EBA: Early Bronze Age; LBA: Late Bronze Age.

We know now that Samic peoples expanded during the Late Iron Age into Palaeo-Laplandic populations, admixing with them and creating this modern cline. Finns expanded later to the north (in one of their known genetic bottlenecks), admixing with (and displacing) the Saami in Finland, especially replacing their male lines.

So how did Ugric and Samoyedic peoples admix with Palaeo-Siberian populations further, to obtain their modern cline? The answer is, logically, with East Asian migrations related to forest-steppe populations of Central Asia after the Mezhovska and Karasuk periods, i.e. during the Iron Age and later. Other groups from the forest-steppe in Central Asia show similar East Asian (“Siberian”) admixture. We know this from Narasimhan et al. (2018):

(…) we observe samples from multiple sites dated to 1700-1500 BCE (Maitan, Kairan, Oy_Dzhaylau and Zevakinsikiy) that derive up to ~25% of their ancestry from a source related to present-day East Asians and the remainder from Steppe_MLBA. A similar ancestry profile became widespread in the region by the Late Bronze Age, as documented by our time transect from Zevakinsikiy and samples from many sites dating to 1500-1000 BCE, and was ubiquitous by the Scytho-Sarmatian period in the Iron Age.

We already have some information about these later migrations:

siberian-genetic-component-chronology
Very important observation with implication of population turnover is that pre-Turkic Inner Eurasian populations’ Siberian ancestry appears predominantly “Uralic-Yeniseian” in contrast to later dominance of “Tungusic-Mongolic” sort (which does sporadically occur earlier). Alexander M. Kim

The Ugric-speaking Sargat culture in Western Siberia shows the expected mixture of haplogroups (ca. 500 BC – 500 AD), with 5 samples of hg N and 2 of hg R1a1, in Pilipenko et al. (2017). Although radiocarbon dates and subclades are lacking, N lineages probably spread late, because of the late and gradual admixture of Siberian cultures into the Sargat melting pot.

The Samoyedic-speaking Tagar culture also shows signs of a genetic turnover in Pilipenko et al. (2018):

The observed reduction in the genetic distance between the Middle Tagar population and other Scythian like populations of Southern Siberia(Fig 5; S4 Table), in our opinion, is primarily associated with an increase in the role of East Eurasian mtDNA lineages in the gene pool (up to nearly half of the gene pool) and a substantial increase in the joint frequency of haplogroups C and D (from 8.7% in the Early Tagar series to 37.5% in the Middle Tagar series). These features are characteristic of many ancient and modern populations of Southern Siberia and adjacent regions of Central Asia, including the Pazyryk population of the Altai Mountains.

Before the Iron Age, the Karasuk and Mezhovska population were probably already somehow ‘to the north’ within the ancient Steppe-Altai cline (see image below9 created by expanding Seima-Turbino- and Andronovo-related populations. During the Iron Age, further Siberian contributions with Iranian expansions must have placed Uralians of the Central Asian forest-steppe areas much closer to today’s Palaeo-Siberian cline.

However, the modern genetic picture was probably fully developed only in historic times, when Samoyedic and Ugric languages expanded to the north, only in part admixing further with Palaeo-Siberian-speaking nomads from the Circum-Arctic region (see here for a recent history of Samoyedic Enets), which justifies their more recent radical ‘northern shift’.

east-uralic-clines
Modified image from Jeong et al. (2018), supplementary materials. 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 north-south 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.

This late acquisition of the language by Palaeo-Siberian nomads (without much population replacement) also justifies the wide PCA clusters of very small Siberian populations. See for example in the PCA from Tambets et al. (2018):

uralic-ugric-samoyedic-modern-clines
Approximate Ugric and Samoyedic clines (exluding apparent outliers). Modified from Tambets et al. (2018). Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations

For their relationship with modern Mansi, we have information on Hungarian conqueror populations from Neparáczki et al. (2018):

Moreover, Y, B and N1a1a1a1a Hg-s have not been detected in Finno-Ugric populations [80–84], implying that the east Eurasian component of the Conquerors and Finno-Ugric people are probably not directly related. The same inference can be drawn from phylogenetic data, as only two Mansi samples appeared in our phylogenetic trees on the side branches (S1 Fig, Networks; 1, 4) suggesting that ancestors of the Mansis separated from Asian ancestors of the Conquerors a long time ago. This inference is also supported by genomic Admixture analysis of Siberian and Northeastern European populations [85], which revealed that Mansis received their eastern Siberian genetic component approximately 5–7 thousand years ago from ancestors of modern Even and Evenki people. Most likely the same explanation applies to the Y-chromosome N-Tat marker which originated from China [86,87] and its subclades are now widespread between various language groups of North Asia and Eastern Europe [88].

The genetic picture of Hungarians (their formed cline with Mansi and their haplogroups) may be quite useful for the true admixture found originally in Mansi peoples at the beginning of the Iron Age. By now it is clear even from modern populations that Steppe_MLBA ancestry accompanied the Uralic expansion to the east (roughly approximated in the graphic with Afanasievo_EBA + Bichon_LP EasternHG_M):

siberian-population-expansions
Admixture modelling using qpAdm. Maps showing locations and ancestry proportions of ancient (left) and modern (right) groups. From Sikora et al. (2018).

Continue reading the final post of the series: Corded Ware—Uralic (IV): Haplogroups R1a and N in Finno-Ugric and Samoyedic.

Related

  • The traditional multilingualism of Siberian populations
  • Iron Age bottleneck of the Proto-Fennic population in Estonia
  • Y-DNA haplogroups of Tuvinian tribes show little effect of the Mongol expansion
  • Corded Ware—Uralic (I): Differences and similarities with Yamna
  • Haplogroup R1a and CWC ancestry predominate in Fennic, Ugric, and Samoyedic groups
  • The Iron Age expansion of Southern Siberian groups and ancestry with Scythians
  • Evolution of Steppe, Neolithic, and Siberian ancestry in Eurasia (ISBA 8, 19th Sep)
  • Mitogenomes from Avar nomadic elite show Inner Asian origin
  • On the origin and spread of haplogroup R1a-Z645 from eastern Europe
  • Oldest N1c1a1a-L392 samples and Siberian ancestry in Bronze Age Fennoscandia
  • Consequences of Damgaard et al. 2018 (III): Proto-Finno-Ugric & Proto-Indo-Iranian in the North Caspian region
  • The concept of “Outlier” in Human Ancestry (III): Late Neolithic samples from the Baltic region and origins of the Corded Ware culture
  • Genetic prehistory of the Baltic Sea region and Y-DNA: Corded Ware and R1a-Z645, Bronze Age and N1c
  • More evidence on the recent arrival of haplogroup N and gradual replacement of R1a lineages in North-Eastern Europe
  • Another hint at the role of Corded Ware peoples in spreading Uralic languages into north-eastern Europe, found in mtDNA analysis of the Finnish population
  • New Ukraine Eneolithic sample from late Sredni Stog, near homeland of the Corded Ware culture
  • Dzudzuana, Sidelkino, and the Caucasus contribution to the Pontic-Caspian steppe

    hunter-gatherer-pottery

    It has been known for a long time that the Caucasus must have hosted many (at least partially) isolated populations, probably helped by geographical boundaries, setting it apart from open Eurasian areas.

    David Reich writes in his book the following about India:

    The genetic data told a clear story. Around a third of Indian groups experienced population bottlenecks as strong or stronger than the ones that occurred among Finns or Ashkenazi Jews. We later confirmed this finding in an even larger dataset that we collected working with Thangaraj: genetic data from more than 250 jati groups spread throughout India (…)

    Rather than an invention of colonialism as Dirks suggested, long-term endogamy as embodied in India today in the institution of caste has been overwhelmingly important for millennia. (…)

    The Han Chinese are truly a large population. They have been mixing freely for thousands of years. In contrast, there are few if any Indian groups that are demographically very large, and the degree of genetic differentiation among Indian jati groups living side by side in the same village is typically two to three times higher than the genetic differentiation between northern and southern Europeans. The truth is that India is composed of a large number of small populations.

    There is little doubt now, based on findings spanning thousands of years, that the Mesolithic and Neolithic Caucasus hosted various very small populations, even if the ancestral components may be reduced to the few known to date (such as ANE, EHG, AME*, ENA, CHG, and other “deep” ancestral components).

    NOTE. I will call the ancestral component of Dzudzuana/Anatolian hunter-gatherers Ancient Middle Easterner (AME), to give a clear idea of its likely extension during the Late Upper Palaeolithic, and to avoid using the more simplistic Dzudzuana, unless it is useful to mention these specific local samples.

    dzudzuana-pca
    Image modified from Lazaridis et al. (2018), including Caucasus, Don-Volga-Ural, and North Pontic Mesolithic-Neolithic populations. “Ancient West Eurasian population structure. (a) Geographical distribution of key ancient West Eurasian populations. (b) Temporal distribution of key ancient West Eurasian populations (approximate date in ky BP). (c) PCA of key ancient West Eurasians, including additional populations (shown with grey shells), in the space of outgroup f4-statistics (Methods).”

    Genetic labs have a strong fixation with ancestry. I guess the use of complex statistical methods gives professionals and laymen alike the feeling of dealing with “Science”, as opposed to academic fields where you have to interpret data. I think language reveals a lot about the way people think, and the fact that ancestral components are called ‘lineages’ – while not wrong per se – is a clear symptom of the lack of interest in the true lineages: Y-DNA haplogroups.

    Y-DNA bottlenecks

    It has become quite clear that male-biased migrations are often the ones which can be confidently followed for actual population movements and ethnolinguistic identification, at least until the Iron Age. The frequently used Palaeolithic clusters offer a clear example of why ancestry does not represent what some people believe: They merely give a basic idea of sizeable population replacements by distant peoples.

    Both concepts are important: sizeable and distant peoples. For example, during the Upper Palaeolithic in Europe there was a sizeable population replacement of the Aurignacian Goyet cluster by the Gravettian Vestonice cluster (probably from populations of far eastern Russia) coupled with the arrival of haplogroup I, although during the thousands of years that this material culture lasted, the previously expanded C1a2 lineages did not disappear, and there were probably different resurgence and admixture events.

    Haplogroup I certainly expanded with the Gravettian culture to Iberia, where the Goyet ancestry did not change much – probably because of male-driven migrations -, to the extent that during the Magdalenian expansions haplogroup I expanded with an ancestry closer to Goyet, in what is called a ‘resurge’ of the Goyet cluster – even though there is a clear replacement of male lines.

    The Villabruna (WHG) cluster is another good example. It probably spread with haplogroup R1b-L754, which – based on the extra ‘East Asian’ affinity of some samples and on modern samples from the Middle East – came probably from the east through a southern route, and not too long before the expansion of WHG likely from around the Black Sea, although this is still unclear. The finding of haplogroup I in samples of mostly WHG ancestry could confuse people that do not care about timing, sub-structured populations, and gene flow.

    palaeolithic-expansions-reich
    Image from David Reich’s Who We Are and How We Got Here. Having migrated out of Africa and the Near East, modern human pioneer populations spread throughout Eurasia (1). By at least thirty-nine thousand years ago, one group founded a lineage of European hunter-gatherers that persisted largely uninterrupted for more than twenty thousand years (2). Eventually, groups derived from an eastern branch of this founding population of European huntergatherers spread west (3), displaced previous groups, and were eventually themselves pushed out of northern Europe by the spread of glacial ice, shown at its maximum extent (top right). As the glaciers receded, western Europe was repeopled from the southwest (4) by a population that had managed to persist for tens of thousands of years and was related to an approximately thirty-five-thousand-year old individual from far western Europe. A later human migration, following the first strong warming period, had an even larger impact, with a spread from the southeast (5) that not only transformed the population of western Europe but also homogenized the populations of Europe and the Near East. At a single site—Goyet Caves in Belgium—ancient DNA from individuals spread over twenty thousand years reflects these transformations, with representatives from the Aurignacian, Gravettian, and Magdalenian periods.

    NOTE. If you don’t understand why ‘clusters’ that span thousands of years don’t really matter for the many Palaeolithic population expansions that certainly happened among hunter-gatherers in Europe, just take a look at what happened with Bell Beakers expanding from Yamna into western Europe within 500 years.

    If we don’t thread carefully when talking about population migrations, these terms are bound to confuse people. Just as the fixation on “steppe ancestry” – which marks the arrival in Chalcolithic Europe of peoples from the Pontic-Caspian region – has confused a lot of researchers to this day.

    When I began to write about the Indo-European demic diffusion model, my concern was to find a single spot where a North-West Indo-European proto-language could have expanded from ca. 2000 BC (our most common guesstimate). Based on the 2015 papers, and in spite of their conclusions, I thought it had become clear that Corded Ware was not it, and it was rather Bell Beakers. I assumed that Uralic was spoken to the north (as was the traditional belief), and thus Corded Ware expanded from the forest zone, hence steppe ancestry would also be found there with other R1a lineages.

    With the publication of Mathieson et al. (2017) and Olalde et al. (2017), I changed my mind, seeing how “steppe ancestry” did in fact appear quite late, hence it was likely to be the result of very specific population movements, probably directly from the Caucasus. Later, Mathieson published in a revision the sample from Alexandria of hg R1a-M417 (probably R1a-Z645, possibly Z93+), which further supported the idea that the migration of Corded Ware peoples started near the North Pontic forest-steppe (as I included in a the next revision).

    The question remains the same I repeated recently, though: where do the extra Caucasus components (i.e. beyond EHG) of Eneolithic Ukraine/Corded Ware and Khvalynsk/Yamna come from?

    Steppe ancestry: “EHG” + “CHG”?

    About EHG ancestry

    From Lazaridis et al. (2018):

    Considering 2-way mixtures, we can model Karelia_HG as deriving 34 ± 2.8% of its ancestry from a Villabruna-related source, with the remainder mainly from ANE represented by the AfontovaGora3 (AG3) sample from Lake Baikal ~17kya.

    AG3 was likely of haplogroup Q1a (as reported by YFull, see Genetiker), and probably the ANE ancestry found in Eastern Europe accompanied a Palaeolithic migration of Q1a2-M25 (formed ca. 22600 BC, TMRCA ca. 14300 BC).

    NOTE. You can read more about the expansion of Q lineages during the Palaeolithic.

    Combined with what we know about the Eneolithic Steppe and Caucasus populations – it is likely that ANE ancestry remained the most important component of some of the small ghost populations of the Caucasus until their emergence with the Lola culture.

    pca-caucasus-dzudzuana
    Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. Marked and labelled are the Balkan samples referenced in this text An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them. See the original file here. To understand the drawn potential Caucasus Mesolithic cluster, see above the PCA from Lazaridis et al. (2018).

    The first sample we have now attributed to the EHG cluster is Sidelkino, from the Samara region (ca. 9300 BC), mtDNA U5a2. In Damgaard et al. (Science 2018), Yamnaya could be modelled as a CHG population related to Kotias Klde (54%) and the remaining from ANE population related to Sidelkino (>46%), with the following split events:

    1. A split event, where the CHG component of Yamnaya splits from KK1. The model inferred this time at 27 kya (though we note the larger models in Sections S2.12.4 and S2.12.5 inferred a more recent split time).
    2. A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
    3. A split event, where the ANE component of Yamnaya splits from Botai. We inferred this to occur 17 kya. Note that this is above the Sidelkino split time, so our model infers Yamnaya to be more closely related to the EHG Sidelkino, as expected.
    4. An ancestral split event between the CHG and ANE ancestral populations. This was inferred to occur around 40 kya.

    Other samples classified as of the EHG cluster:

    • Popovo2 (ca. 6250 BC) of hg J1, mtDNA U4d – Po2 and Po4 from the same site (ca. 6550 BC) show continuity of mtDNA.
    • Karelia_HG, from Juzhnii Oleni Ostrov (ca. 6300 BC): I0211/UzOO40 (ca. 6300 BC) of hg J1(xJ1a), mtDNA U4a; and I0061/UzOO74 of hg R1a1(xR1a1a), mtDNA C1
    • UzOO77 and UzOO76 from Juzhnii Oleni Ostrov (ca. 5250 BC) of mtDNA R1b.
    • Samara_HG from Lebyanzhinka (ca. 5600 BC) of hg R1b1a, mtDNA U5a1d.

    From the analysis of Lazaridis et al. (2018), we have some details about their admixture:

    dzudzuana-admixture-sidelkino
    Image modified from Lazaridis et al. (2018). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (Left) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown. (Right) ‘Speculative’ estimates. The highest number of sources (≤5) with admixture estimates within [0,1] are shown for each population. Some of the admixture proportions are not significantly different from 0 (Supplementary Information section 4).

    About Anatolia_Neolithic ancestry

    About the enigmatic Anatolia_Neolithic-related ancestry found in Pontic-Caspian steppe samples, this is what Wang et al. (2018) had to say:

    We focused on model of mixture of proximal sources such as CHG and Anatolian Chalcolithic for all six groups of the Caucasus cluster (Eneolithic Caucasus, Maykop and Late Makyop, Maykop-Novosvobodnaya, Kura-Araxes, and Dolmen LBA), with admixture proportions on a genetic cline of 40-72% Anatolian Chalcolithic related and 28-60% CHG related (Supplementary Table 7). When we explored Romania_EN and Greece_Neolithic individuals as alternative southeast European sources (30-46% and 36-49%), the CHG proportions increased to 54-70% and 51-64%, respectively. We hypothesize that alternative models, replacing the Anatolian Chalcolithic individual with yet unsampled populations from eastern Anatolia, South Caucasus or northern Mesopotamia, would probably also provide a fit to the data from some of the tested Caucasus groups.

    Also:

    The first appearance of ‘Near Eastern farmer related ancestry’ in the steppe zone is evident in Steppe Maykop outliers. However, PCA results also suggest that Yamnaya and later groups of the West Eurasian steppe carry some farmer related ancestry as they are slightly shifted towards ‘European Neolithic groups’ in PC2 (Fig. 2D) compared to Eneolithic steppe. This is not the case for the preceding Eneolithic steppe individuals. The tilting cline is also confirmed by admixture f3-statistics, which provide statistically negative values for AG3 as one source and any Anatolian Neolithic related group as a second source

    yamnaya-caucasus-dzudzuana
    Modified image from Wang et al. (2018). In blue, Yamna-related populations. In red, Corded Ware-related populations, and two elevated Anatolia_Neolithic values in Yamna. Notice how only GAC-related admixture increases the Anatolian_N-related ancestry in the Yamna outlier from Ozero, and the late Yamna sample from Hungary, related to the homogeneous Yamna population. “Supplementary Table 14. P values of rank=3 and admixture proportions in modelling Steppe ancestry populations as a four-way admixture of distal sources EHG, CHG, Anatolian_Neolithic and WHG using 14 outgroups.Left populations: Steppe cluster, EHG, CHG, WHG, Anatolian_Neolithic. Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.”

    Detailed exploration via D-statistics in the form of D(EHG, steppe group; X, Mbuti) and D(Samara_Eneolithic, steppe group; X, Mbuti) show significantly negative D values for most of the steppe groups when X is a member of the Caucasus cluster or one of the Levant/Anatolia farmer-related groups (Supplementary Figs. 5 and 6). In addition, we used f- and D-statistics to explore the shared ancestry with Anatolian Neolithic as well as the reciprocal relationship between Anatolian- and Iranian farmer-related ancestry for all groups of our two main clusters and relevant adjacent regions (Supplementary Fig. 4). Here, we observe an increase in farmer-related ancestry (both Anatolian and Iranian) in our Steppe cluster, ranging from Eneolithic steppe to later groups. In Middle/Late Bronze Age groups especially to the north and east we observe a further increase of Anatolian farmer related ancestry consistent with previous studies of the Poltavka, Andronovo, Srubnaya and Sintashta groups and reflecting a different process not especially related to events in the Caucasus.

    (…) Surprisingly, we found that a minimum of four streams of ancestry is needed to explain all eleven steppe ancestry groups tested, including previously published ones (Fig. 2; Supplementary Table 12). Importantly, our results show a subtle contribution of both Anatolian farmer-related ancestry and WHG-related ancestry (Fig.4; Supplementary Tables 13 and 14), which was likely contributed through Middle and Late Neolithic farming groups from adjacent regions in the West. The discovery of a quite old AME ancestry has rendered this probably unnecessary, because this admixture from an Anatolian-like ghost population could be driven even by small populations from the Caucasus.

    yamna-caucasus-cwc-anatolia-neolithic
    Image modified from Wang et al. (2018). Marked are: in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus 1128 cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups (see also Supplementary Tables 10, 14 and 20).”

    NOTE. For a detailed account of the possibilities regarding this differential admixture in the North Pontic area in contrast to the Don-Volga-Ural region, you can read the posts Sredni Stog, Proto-Corded Ware, and their “steppe admixture”, and Corded Ware culture origins: The Final Frontier.

    While it is not yet fully clear, the increased Anatolian_Neolithic-like ancestry in Ukraine_Eneolithic samples (see below) makes it unlikely that all such ancestry in Corded Ware groups comes from a GAC-related contribution. It is likely that at least part of it represents contributions from populations of the Caucasus, based on the mostly westward population movements in the steppe from ca. 4600 BC on, including the Suvorovo-Novodanilovka expansion, and especially the Kuban-Maykop expansion during the final Eneolithic into the North Pontic area.

    NOTE. Since CHG-like groups from the Caucasus may have combinations of AME and ANE ancestry similar to Yamna (which may thus appear as ‘steppe ancestry’ in the North Pontic area), it is impossible to interpret with precision the following ADMIXTURE graphic:

    ukraine-whg-ehg-steppe
    Modified image from Mathieson et al. (2018). Supervised ADMIXTURE analysis, modelling each ancient individual (one per row) as a mixture of population clusters constrained to contain northwestern-Anatolian Neolithic (grey), Yamnaya from Samara (yellow), EHG (pink) and WHG (green) populations. Dates in parentheses indicate approximate range of individuals in each population.

    North-Eastern Technocomplex

    The East Asian contribution to samples from the WHG samples (like Loschbour or La Braña), as specified in Fu et al. (2016), does not seem to be related to Baikal_EN, and appears possibly (in the ADMIXTURE analysis) integrated into he Villabruna component. I guess this implies that the shared alleles with East Asians are quite early, and potentially due to the expansion of R1b-L754 from the East.

    It would be interesting to know the specific material culture Sidelkino belonged to – i.e. if it was related to the expansion of the North-Eastern Technocomplex – , and its Y-DNA. The Post-Swiderian expansion into eastern Europe, probably associated with the expansion of R1b-P297 lineages (including R1b-M73, found later in Botai and in Baltic HG) is supposed to have begun during the 11th millennium BC, but migrations to the Urals and beyond are probably concentrated in the 9th millennium, so this sample is possibly slightly early for R1b.

    NOTE. User Rozenfeld at Anthrogenica posted this, which I think is interesting (in case anyone wants to try a Y-SNP call):

    there is something strange with Sidelkino EHG: first, its archaeological context is not described in the supplementary. Second, its sex is not listed in the supplementary tables. Third, after looking for info about this sample, I found that: “Сиделькино-3. Для снятия вопроса о половой принадлежности индивида была проведена генетическая экспертиза, выявившая принадлежность останков мужчине.”(translation: Sidelkino-3. To resolve the question about sex of the remains, the genetic analysis was conducted, which showed that remains belonged to male), source: http://static.iea.ras.ru/books/7487_Traditsii.pdf

    So either they haven’t mentioned his Y-DNA in the paper for some reason, or there are more than one Sidelkino sample and the male one has not yet been published. The coverage of the Sidelkino sample from the paper is 2.9, more than enough to tell Y-DNA haplogroup.

    zaliznyak-post-swiderian
    The map of spreading of Post-Swiderian and Post-Krasnosillian sites in Mesolithic of Eastern Europe in the 8th millennia BC. From Zaliznyak (see here).

    My speculative guess right now about specific population movements in far eastern Europe, based on the few data we have:

    • The expansion of the North-Eastern Technocomplex first around the 9th millennium BC, most likely expanded R1b-P279 ca. 11300 BC, judging by its TMRCA, with both R1b-M73 (TMRCA 5300) and R1b-M269 (TMRCA 4400 BC) info (with extra El Mirón ancestry) back, and thus Eurasiatic.
    • The expansion of haplogroup J1 to the north may have happened before or after the R1b-P279 expansion. Judging by the increase in AG3-related ancestry near Karelia compared to Baltic_HG, it is possible that it expanded just after R1b-P279 (hence possibly J1-Y6304? TMRCA 9700 BC). Its long-lasting presence in the Caucasus is supported by the Satsurblia (ca. 11300 BC) and the Dolmen BA (ca. 1300 BC) samples.
    • The expansion of R1a-M17 ca. 6600 BC is still likely to have happened from the east, based on the R1a-M17 samples found in Baikalic cultures slightly later (ca. 5300 BC). The presence of elevated Baikal_EN ancestry in Karelia HG and in Samara HG, and the finding of R1a-M417 samples in the Forest Zone after the Mesolithic suggests a connection with the expansion of Hunter-Gatherer pottery, from the Elshanka culture in the Samara region northward into the Forset Zone and westward into the North Pontic area.
    • The expansion of R1b-M73 ca. 5300 BC is likely to be associated with the emergence of a group east of the Urals (related to the later Botai culture, and potentially Pre-Yukaghir). Its presence in a Narva sample from Donkalnis (ca. 5200 BC) suggest either an early split and spread of both R1b-P297 lineages (M73 and M269) through Eastern Europe, or maybe a back-migration with hunter-gatherer pottery.
    • R1b-M269 spread successfully ca. 4400 BC (and R1b-L23 ca. 4100 BC, both based on TMRCA), and this successful expansion is probably to be associated with the Khvalynsk-Novodanilovka expansion. We already know that Samara_HG ca. 5600 was R1b1a, so it is likely that R1b-M269 appeared (or ‘resurged’) in the Volga-Ural region shortly after the expansion of R1a-M17, whose expansion through the region may be inferred by the additional AG3 and Baikal_EN ancestry. Interesting from Samara_HG compared to the previous Sidelkino sample is the introduction of more El Mirón-related ancestry, typical of WHG populations (and thus proper of Baltic groups).

    NOTE. The TMRCA dates are obviously gross approximations, because a) the actual rate of mutation is unknown and b) TMRCA estimates are based on the convergence of lineages that survived. The potential finding of R1a-Z645 (possibly Z93+) in Ukraine Eneolithic (ca. 4000 BC), and the potential finding of R1b-L23 in Khvalynsk ca. 4250 BC complicates things further, in terms of dates and origins of any subclade.

    The question thus remains as it was long ago: did R1b-M269 lineages expand (‘return’) from the east, near the Urals, or directly from the north? Were they already near Samara at the same time as the expansion of hunter-gatherer pottery, and were not much affected by it? Or did they ‘resurge’ from populations admixed with Caucasus-related ancestry after the expansion of R1a-M17 with this pottery (since there are different stepped expansions from the Samara region)? We could even ask, did R1a-M17 really expand from the east, i.e. are the dates on Baikalic subclades from Moussa et al. (2016) reliable? Or did R1a-M17 expand from some pockets in the Pontic-Caspian steppe, taking over the expansion of HG pottery at some point?

    hunger-gatherer-pottery
    Early Neolithic cultures in eastern and central Europe: 1–Yelshanian; 2–North Caspian; 3–Rakushechnyj Yar; 4–Surskian; 5–Dnieper-Donetsian; 6– Bug-Dniesterian; 7–Upper Volga; 8–Narvian; 9–Linear Pottery. White arrows: expansion of early farming; black arrows: spread of pottery-making traditions. From Dolukhanov et al. (2009).

    Maglemose-related migrations

    The most interesting aspect from the new paper (regarding Indo-Uralic migrations) is that Ancestral Middle Easterner ancestry will probably be a better proxy for the Anatolia_Neolithic component found in Ukraine Mesolithic to Eneolithic, and possibly also for some of the “more CHG-like” component found among Pontic-Caspian steppe populations, all likely derived from different admixture events with groups from the Caucasus.

    NOTE. Even the supposed gene flow of Neolithic Iranian ancestry into the Caucasus can be put into question, since that means possibly a Dzudzuana-like population with greater “deep ancestry” proportion than the one found in CHG, which may still be found within the Caucasus.

    If it was not clear already that following ‘steppe ancestry’ wherever it appears is a rather lame way of following Indo-European migrations, every single sample from the Caucasus and their admixture with Pontic-Caspian steppe populations will probably show that “steppe ancestry” is in fact formed by a variety of steppe-related ancestral components, impossible to follow coherently with a single population. Exactly what is happening already with the Siberian ancestry.

    If the paper on the Dzudzuana samples has shown something, is that the expansion of an ANE-like population shook the entire Caucasus area up to the Zagros Mountains, creating this ANE – AME cline that are CHG and Iran_N, with further contributions of “deep ancestries” (probably from the south) complicating the picture further.

    If this happens with few known samples, and we know of an ANE-like ghost population in the Caucasus (appearing later in the Lola culture), we can already guess that the often repeated “CHG component” found in Ukraine_Eneolithic and Khvalynsk will not be the same (except the part mediated by the Novodanilovka expansion).

    This ANE-like expansion happened probably in the Late Upper Palaeolithic, and reached Northern Europe probably after the expansion of the Villabruna cluster (ca. 12000 BC), judging by the advance of AG3-like and ENA-like ancestry in later WHG samples.

    The population movements during the Mesolithic and Early Neolithic in the North Pontic area are quite complicated: the extra AME ancestry is probably connected to the admixture with populations from the Caucasus, while the close similarity of Ukraine populations with Scandinavian ones (with an increase in Villabruna ancestry from Mesolithic to Neolithic samples), probably reveal population movements related to the expansion of Maglemose-related groups.

    maglemose-mesolithic
    Etno-cultural situation in Central and Eastern Europe in the Late Mesolithic — Early Neolithic (VI—V Mill. BC) (after Конча 2004: 201, карта 1; made after ideas by L. L. Zaliznyak). Legend: 1 — Maglemose circle in the VII Mill. BC (after Gr. Clark); 2—7 — Mesolithic cultures of the Post-Maglemose tradition, VI Mill. BC (after S. Kozłowsky, L. L. Zaliznyak): 2 — de Leyen-Wartena; 3 — Oldesloe — Godenaa; 4 — Chojnice — Peńki; 5 — Janisłavice; 6 — finds of Janisłavice artefacts outside of the main area; 7 — Donets culture; 8 — directions of the settling of Janisłavice people (after S. Kozłowsky and L. L. Zaliznyak); 9 — the south border of Mesolithic and Early Neolithic cultures of post-Swidrian and post-Arensburgian traditions; 10 — northern border of settlement of the Balkan-Danubian farmers; 11 — Bug- Dniester culture; 12 — Neolithic cultures emerged on the ethno-cultural basis of post-Maglemose: Э — Ertebölle-Ellerbeck, Н — Neman, Д — Dnieper-Donets, М — Mariupol (western variants). From Klein (2017).

    These Maglemose-related groups were probably migrants from the north-west, originally from the Northern European Plains, who occupied the previous Swiderian territory, and then expanded into the North Pontic area. The overwhelming presence of I2a (likely all I2a2a1b1b) lineages in Ukraine Neolithic supports this migration.

    The likely picture of Mesolithic-Neolithic migrations in the North Pontic area right now is then:

    1. Expansion of R1a-M459 from the east ca. 12000 BC – probably coupled with AG3 and also some Baikal_EN ancestry. First sample is I1819 from Vasilievka (ca. 8700 BC), another is from Dereivka ca. 6900 BC.
    2. Expansion of R1b-V88 from the Balkans in the west ca. 9700 BC, based on its TMRCA and also the Balkan hunter-gatherer population overwhemingly of this haplogroup from the 10th millennium until the Neolithic. First sample is I1734 from Vasilievka (ca. 7252 BC), which suggests that it replaced the male population there, based on their similar EHG-like adxmixture (and lack of sizeable WHG increase), and shared mtDNA U5b2, U5a2.
    3. Expansion of I2a-Y5606 probably ca. 6800 based on its TMRCA with Janislawice culture. Supporting this is the increase in WHG contribution to Neolithic samples, including the spread of U4 subclades compared to the previous period.
    4. Expansion of R1a-M17 starting probably ca. 6600 BC in the east (see above).

    NOTE. The first sample of haplogroup I appears in the Mesolithic: I1763 (ca. 8100 BC) of haplogroup I2a1, probably related to an older Upper Palaeolithic expansion.

    janislawice
    Distribution of archeological cultures in the North Pontic Region during the Mesolithic (7th – 6th millennium BCE). Dotted, dashed and solid lines with corresponding arrows indicate alternative models of the spread of the Grebenyky culture groups. (After Bryuako IV., Samojlova TL., Eds, Drevnie kul’tury Severo-­‐Zapadnogo Prichernomor’ya, Odessa: SMIL, 2013.) Nikitin – Ivanova 2017.

    Conclusion

    It is becoming more and more clear with each new paper that – unless the number of very ancient samples increases – the use of Y-chromosome haplogroups remains one of the most important tools for academics; this is especially so in the steppes, in light of the diversity found in populations from the Caucasus. A clear example comes from the Yamna – Corded Ware similarities:

    After the publication of the 2015 papers, it was likely that Yamna expanded with haplogroup R1b-L23, but it has only become crystal clear that Yamna expanded through the steppes into Bell Beakers, now that we have data about the strict genetic homogeneity of the whole Yamna population from west to east (including Afanasevo), in contrast with contemporary Corded Ware peoples which expanded from a different forest-steppe population.

    The presence of haplogroups Q and R1a-M459 (xM17) in Khvalynsk along with a R1b1a sample, which some interpreted as being akin to modern ‘mixed’ populations in the past, is likely to point instead to a period of Khvalynsk-Novodanilovka expansion with R1b-M269, where different small populations from the steppe were being integrated into the common Khvalynsk stock, but where differences are seen in material culture surrounding their burials, as supported by the finding of R1b1 in the Kuban area already in the first half of the 5th millennium. The case would be similar to the early ‘mixed’ Icelandic population.

    Only after the emergence of the Samara culture (in the second half of the 6th millennium BC), with a sample of haplogroup R1b1a, starts then the obvious connection with Early Proto-Indo-Europeans; and only after the appearance of late Sredni Stog and haplogroup R1a-M417 (ca. 4000 BC) is its connection with Uralic also clear. In previous population movements, I think more haplogroups were involved in migrations of small groups, and only some communities among them were eventually successful, expanding to be dominant, creating ever growing cultures during their expansions.

    Indeed, if you think in terms of Uralic and Indo-European just as converging languages, and forget their potential genetic connection, then the genetic + linguistic picture becomes simplified, and the upper frontier of the 6th millennium BC with a division North Pontic (Mariupol) vs. Volga-Ural (Samara) is enough. However, tracing their movements backwards – with cultural expansions from west to east (with the expansion of farming), and earlier east to west (with hunter-gatherer pottery), and still earlier west to east (with the north-eastern technocomplex), offers an interesting way to prove their potential connection to macrofamilies, at least in terms of population movements.

    corded-ware-uralic-qpgraph
    Modified image from Tambets et al. (2018) Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. a The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. The NeolL (Neolithic Levant) ancestry selected in this qpGraph is likely to correspond (at least in part) to a specific Dzudzuana-like component present in the CHG-like population that admixed in the North Pontic area.

    I am quite convinced right now that it would be possible to connect the expansion of R1b-L754 subclades with a speculative Nostratic (given the R1b-V88 connection with Afroasiatic, and the obvious connection of R1b-L297 with Eurasiatic). Paradoxically, the connection of an Indo-Uralic community in the steppes (after the separation of Yukaghir) with any lineage expansion (R1a-M17, R1b-M269, or even Q, I or J1) seems somehow blurrier than one year ago, possibly just because there are too many open possibilities.

    David Reich says about the admixture with Neanderthals, which he helped discover:

    At the conclusion of the Neanderthal genome project, I am still amazed by the surprises we encountered. Having found the first evidence of interbreeding between Neanderthals and modern humans, I continue to have nightmares that the finding is some kind of mistake. But the data are sternly consistent: the evidence for Neanderthal interbreeding turns out to be everywhere. As we continue to do genetic work, we keep encountering more and more patterns that reflect the extraordinary impact this interbreeding has had on the genomes of people living today.

    I think this is a shared feeling among many of us who have made proposals about anything, to fear that we have made a gross, evident mistake, and constantly look for flaws. However, it seems to me that geneticists are more preoccupied with being wrong in their developed statistical methods, in the theoretical models they are creating, and not so much about errors in the true ancient ethnolinguistic picture human population genetics is (at least in theory) concerned about. Their publications are, after all, constantly associating genetic finds with cultures and (whenever possible) languages, so this aspect of their research should not be taken lightly.

    Seeing how David Anthony or Razib Khan (among many others) have changed their previously preferred migration models as new data was published, and they continue to be respected in their own fields, I guess we can be confident that professionals with integrity are going to accept whatever new picture appears. While I don’t think that genetic finds can change what we can reconstruct with comparative grammar, I am also ready to revise guesstimates and routes of expansion of certain dialects if R1a-Z645 is shown to have accompanied Late Proto-Indo-Europeans during their expansion with Yamna, and later integrated somehow with Corded Ware.

    However, taking into account the obsession of some with an ancestral, uninterrupted R1a—Indo-European association, and the lack of actual political repercussion of Neanderthal admixture, I think the most common nightmare that all genetic researchers should be worried about is to keep inflating this “Yamnaya ancestry”-based hornet’s nest, which has been constantly stirred up for the past two years, by rejecting it – or, rather, specifying it into its true complex nature.

    This succession of corrections and redefinitions, coupled with the distinct Y-DNA bottleneck of each steppe population, will eventually lead to a completely different ethnolinguistic picture of the Pontic-Caspian region during the Eneolithic, which is likely to eventually piss off not only reasonable academics stubbornly attached to the CWC-IE idea, but also a part of those interested in daydreaming about their patrilineal ancestors.

    Sometimes it’s better to just rip off the band-aid once and for all…

    Featured image from The oldest pottery in hunter-gatherer communitiesand models of Neolithisation of Eastern Europe (2015), by Andrey Mazurkevich and Ekaterina Dolbunova.

    Related

    Interesting is today’s post in Ancient DNA Era: Is Male-driven Genetic Replacement always meaning Language-shift?

    Neolithic and Bronze Age Anatolia, Urals, Fennoscandia, Italy, and Hungary (ISBA 8, 20th Sep)

    jena-isba8

    I will post information on ISBA 8 sesions today as I see them on Twitter (see programme in PDF, and sessions from yesterday).

    Official abstracts are listed first (emphasis mine), then reports and images and/or link to tweets. Here is the list for quick access:

    Russian colonization in Yakutia

    Exploring the genomic impact of colonization in north-eastern Siberia, by Seguin-Orlando et al.

    Yakutia is the coldest region in the northern hemisphere, with winter record temperatures below minus 70°C. The ability of Yakut people to adapt both culturally and biologically to extremely cold temperatures has been key to their subsistence. They are believed to descend from an ancestral population, which left its original homeland in the Lake Baykal area following the Mongol expansion between the 13th and 15th centuries AD. They originally developed a semi-nomadic lifestyle, based on horse and cattle breeding, providing transportation, primary clothing material, meat, and milk. The early colonization by Russians in the first half of the 17th century AD, and their further expansion, have massively impacted indigenous populations. It led not only to massive epidemiological outbreaks, but also to an important dietary shift increasingly relying on carbohydrate-rich resources, and a profound lifestyle transition with the gradual conversion from Shamanism to Christianity and the establishment of new marriage customs. Leveraging an exceptional archaeological collection of more than a hundred of bodies excavated by MAFSO (Mission Archéologique Française en Sibérie Orientale) over the last 15 years and naturally kept frozen by the extreme cold temperatures of Yakutia, we have started to characterize the (epi)genome of indigenous individuals who lived from the 16th to the 20th century AD. Current data include the genome sequence of approximately 50 individuals that lived prior to and after Russian contact, at a coverage from 2 to 40 fold. Combined with data from archaeology and physical anthropology, as well as microbial DNA preserved in the specimens, our unique dataset is aimed at assessing the biological consequences of the social and biological changes undergone by the Yakut people following their neolithisation by Russian colons.

    NOTE: For another interesting study on Yakutian tribes, see Relationships between clans and genetic kin explain cultural similarities over vast distances.

    Ancient DNA from a Medieval trading centre in Northern Finland

    Using ancient DNA to identify the ancestry of individuals from a Medieval trading centre in Northern Finland, by Simoes et al.

    Analyzing genomic information from archaeological human remains has proved to be a powerful approach to understand human history. For the archaeological site of Ii Hamina, ancient DNA can be used to infer the ancestries of individuals buried there. Situated approximately 30 km from Oulu, in Northern Finland, Ii Hamina was an important trade place since Medieval times. The historical context indicates that the site could have been a melting pot for different cultures and people of diversified genetic backgrounds. Archaeological and osteological evidence from different individuals suggest a rich diversity. For example, stable isotope analyses indicate that freshwater and marine fish was the dominant protein source for this population. However, one individual proved to be an outlier, with a diet containing relatively more terrestrial meat or vegetables. The variety of artefacts that was found associated with several human remains also points to potential differences in religious beliefs or social status. In this study, we aimed to investigate if such variation could be attributed to different genetic ancestries. Ten of the individuals buried in Ii Hamina’s churchyard, dating to between the 15th and 17th century AD, were screened for presence of authentic ancient DNA. We retrieved genome-wide data for six of the individuals and performed downstream analysis. Data authenticity was confirmed by DNA damage patterns and low estimates of mitochondrial contamination. The relatively recent age of these human remains allows for a direct comparison to modern populations. A combination of population genetics methods was undertaken to characterize their genetic structure, and identify potential familiar relationships. We found a high diversity of mitochondrial lineages at the site. In spite of the putatively distant origin of some of the artifacts, most individuals shared a higher affinity to the present-day Finnish or Late Settlement Finnish populations. Interestingly, different methods consistently suggested that the individual with outlier isotopic values had a different genetic origin, being more closely related to reindeer herding Saami. Here we show how data from different sources, such as stable isotopes, can be intersected with ancient DNA in order to get a more comprehensive understanding of the human past.

    A closer look at the bottom left corner of the poster (the left columns are probably the new samples):

    finland-medieval-admixture

    Plant resources processed in HG pottery from the Upper Volga

    Multiple criteria for the detection of plant resources processed in hunter-gatherer pottery vessels from the Upper Volga, Russia, by Bondetti et al.

    In Northern Eurasia, the Neolithic is marked by the adoption of pottery by hunter-gatherer communities. The degree to which this is related to wider social and lifestyle changes is subject to ongoing debate and the focus of a new research programme. The use and function of early pottery by pre-agricultural societies during the 7th-5th millennia BC is of central interest to this debate. Organic residue analysis provides important information about pottery use. This approach relies on the identification and isotopic characteristics of lipid biomarkers, absorbed into the pores of the ceramic or charred deposits adhering to pottery vessel surfaces, using a combined methodology, namely GC-MS, GC-c-IRMS and EA-IRMS. However, while animal products (e.g., marine, freshwater, ruminant, porcine) have the benefit of being lipid-rich and well-characterised at the molecular and isotopic level, the identification of plant resources still suffers from a lack of specific criteria for identification. In huntergatherer contexts this problem is exacerbated by the wide range of wild, foraged plant resources that may have been potentially exploited. Here we evaluate approaches for the characterisation of terrestrial plant food in pottery through the study of pottery assemblages from Zamostje 2 and Sakhtysh 2a, two hunter-gatherer settlements located in the Upper Volga region of Russia.

    GC-MS analysis of the lipids, extracted from the ceramics and charred residues by acidified methanol, suggests that pottery use was primarily oriented towards terrestrial and aquatic animal products. However, while many of the Early Neolithic vessels contain lipids distinctive of freshwater resources, triterpenoids are also present in high abundance suggesting mixing with plant products. When considering the isotopic criteria, we suggest that plants were a major commodity processed in pottery at this time. This is supported by the microscopic identification of Viburnum (Viburnum Opulus L.) berries in the charred deposits on several vessels from Zamostje.

    The study of Upper Volga pottery demonstrated the importance of using a multidisciplinary approach to determine the presence of plant resources in vessels. Furthermore, this informs the selection of samples, often subject to freshwater reservoir effects, for 14C dating.

    Studies on hunter-gatherer pottery – appearing in eastern Europe before Middle Eastern Neolithic pottery – may be important to understand the arrival of R1a-M17 lineages to the region before ca. 7000 BC. Or not, right now it is not very clear what happened with R1b-P297 and R1a-M17, and with WHG—EHG—ANE ancestry

    Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe

    Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, by Warinner et al.

    Recent paleogenomic studies have shown that migrations of Western steppe herders (WSH), beginning in the Eneolithic (ca. 3300-2700 BCE), profoundly transformed the genes and cultures of Europe and Central Asia. Compared to Europe, the eastern extent of this WSH expansion is not well defined. Here we present genomic and proteomic data from 22 directly dated Bronze Age khirigsuur burials from Khövsgöl, Mongolia (ca. 1380-975 BCE). Only one individual showed evidence of WSH ancestry, despite the presence of WSH populations in the nearby Altai-Sayan region for more than a millennium. At the same time, LCMS/ MS analysis of dental calculus provides direct protein evidence of milk consumption from Western domesticated livestock in 7 of 9 individuals. Our results show that dairy pastoralism was adopted by Bronze Age Mongolians despite minimal genetic exchange with Western steppe herders.

    Detail of the images:

    mongol-bronze-age-pca

    mongol-bronze-age-f4-ancestry

    The origin of social complexity in the development of the Sintashta culture

    kamenni-ambar

    Very interesting PhD thesis by Igor Chechushkov, Bronze Age human communities in the Southern Urals steppe: Sintashta-Petrovka social and subsistence organization (2018).

    Abstract:

    Why and how exactly social complexity develops through time from small-scale groups to the level of large and complex institutions is an essential social science question. Through studying the Late Bronze Age Sintashta-Petrovka chiefdoms of the southern Urals (cal. 2050–1750 BC), this research aims to contribute to an understanding of variation in the organization of local communities in chiefdoms. It set out to document a segment of the Sintashta-Petrovka population not previously recognized in the archaeological record and learn about how this segment of the population related to the rest of the society. The Sintashta-Petrovka development provides a comparative case study of a pastoral society divided into sedentary and mobile segments.

    Subsurface testing on the peripheries of three Sintashta-Petrovka communities suggests that a group of mobile herders lived outside the walls of the nucleated villages on a seasonal basis. During the summer, this group moved away from the village to pasture livestock farther off in the valley, and during the winter returned to shelter adjacent to the settlement. This finding illuminates the functioning of the year-round settlements as centers of production during the summer so as to provide for herd maintenance and breeding and winter shelter against harsh environmental conditions.

    The question of why individuals chose in this context to form mutually dependent relationships with other families and thus give up some of their independence can be answered with a combination of two necessities: to remain a community in a newly settled ecological niche and to protect animals from environmental risk and theft. Those who were skillful at managing communal construction of walled villages and protecting people from military threats became the most prominent members of the society. These people formed the core of the chiefdoms but were not able to accumulate much wealth and other possessions. Instead, they acquired high social prestige that could even be transferred to their children. However, this set of relationships did not last longer than 300 years. Once occupation of the region was well established the need for functions served by elites disappeared, and centralized chiefly communities disintegrated into smaller unfortified villages.

    sintashta-petrovka-archaeological
    Research area: map of the Sintashta-Petrovka archaeological sites. Settlements: 101 – Stepnoye; 102 – Shibaeyvo 1; 103 – Chernorechye 3; 104 – Bakhta; 105 – Paris; 106 – Isiney; 107 – Kuisak; 108 – Ust’ye; 109 – Rodniki; 110 – Konoplyanka; 111 – Zhurumbay; 112 – Arkaim; 113 – Sintashta; 114 – Sintashta 2; 115 – Kamennyi Ambar; 116 – Alandskoye; 117 – Chekatay; 118 – Selek; 119 – Sarym- Sakly; 120 – Kamysty; 121 – Kizilskoye; 122 – Bersuat; 123 – Andreyevskoe; 124 – Ulak; 125 – Streletskoye; 126 – Zarechnoye 4; 127 – Kamennyi Brod. Cemeteries: 201 – Ozernoye 1; 202 – Krivoe Ozero; 203 – Stepnoye M; 204 – Kamennyi Ambar-5; 205 – Stepnoye 1; 206 – Tsarev Kurgan; 207 – Ubagan 2; 208 – Solntse 2; 209 – Bolshekaraganskyi; 210 – Aleksandrovsky 4; 211 – Sintashta; 212 – Solonchanka 1a; 213 – Knyazhenskyi; 214 – Bestamak; 215 – Ishkinovka 1; 216 – Ishkinovka 2; 217 – Novo–Kumakskyi; 218 – Zhaman–Kargala 1; 219 – Tanabergen 2; 220 – Novo-Petrovka; 221 – Semiozernoye 2; 222 – Khalvayi 3

    Some interesting excerpts (emphasis mine):

    The quintessential archaeological evidence of Sintashta-Petrovka communities takes the form of highly nucleated and fortified settlements paired with easily-recognized kurgan (burial mound) cemeteries. This pattern spread across Northern Central Eurasia in a relatively short period of about 300 years (cal. 2050–1750 BC), and the period consists of two chronological phases (Hanks et al. 2007). The earlier Sintashta phase (cal. 2050–1850 BC) is distinguished from the later Petrovka phase (cal. 1850–1750 BC) by some differences in ceramic styles and some techniques of bronze metallurgy (Degtyareva et al. 2001; Vinogradov 2013). Bronze Age subsistence patterns apparently relied on a wide variety of resources, among which meat and milk production played a major role (…). The most outstanding graves are individual male burials accompanied by weaponry (projectile weapons and chariots), the insignia of power (stone mace heads), craft tools, and a specific set of sacrificed animals (horses, cows, and dogs). (…) there were at least two adults buried with chariots and one with sacrificed horses (Epimakhov 1996b). Chariots – the most famous and spectacular material component of Sintashta-Petrovka society – are known exclusively from burial contexts. Two-wheeled vehicles represent complex technology, incorporating some crucial innovations and the investment of substantial resources. Highly developed craft and military skills were required for their production and use. Burials with chariots probably represent military elites who used them (Anthony 2009; Chechushkov 2011; Frachetti 2012:17) and played especially important social roles in Sintashta-Petrovka societies. This pattern strongly suggests that military leadership extended into the realm of ideology and general social prestige (Earle 2011:32–33).

    The following sequence of archaeological cultures – based on the sample of radiocarbon dates (Epimakhov 2007a; 2010a), – is adopted: (1) the Sintashta-Petrovka phase 1 dated to cal. 2050–1750 BC and (2) the Srubnaya-Alakul’ phase 2 dated to cal. 1750–1350 BC.

    (…) control of craft might have provided a source of power for elites in the fortified settlements (Steponaitis 1991). Some bronze tools, such as chisels, adzes, and handsaws seem more abundantly represented at some fortified settlements than at others, raising the possibility of a stronger focus on different craft products and some degree of exchange and interdependence between fortified settlements. (…) Zdanovich (1995:35) estimates 2500 people within the walls at Arkaim. He bases his conclusion an average house size of 140 m2 and the idea that Arkaim households consisted of an extended family of several generations, similar to Iroquois longhouse inhabitants. He also suggests that the entire population did not live in the “town” all the time, but moved around. The fully permanent residents were shamans, warriors, and craftsmen, i.e., elites and attached specialists.

    Summarizing, excavated households represent very strongly similar architectural patterns, similar levels of wealth and prestige, little productive differentiation, and no evidence of elites amassing wealth through control of craft or subsistence production or any other mechanism (Earle 1987). These observations sharply contradict the burial record, where strong social differentiation is visible. The description above recalls the Regional Classic period elites of the Alto Magdalena whose standard of living differed little if at all from anyone else’s. Their elaborate tombs and sculptures suggest supernatural powers and ritual roles were much more important bases of their social prominence than economic control or accumulation of wealth (Drennan 1995:96–97). On the other hand, craft activities (especially metal production) are highly obvious in the Sintashta-Petrovka settlements. Defensive functions could also have played some role for the entire population. This benefit might attract people in an unstable or wild environment to spend much of their time in or near such settlements (Earle 2011:32–33). Since the construction of ditches and outer walls, as well as dwellings with shared walls, requires planning and organization, purposeful collective effort must have been a key feature of Sintashta-Petrovka communities (Vinogradov 2013; Zdanovich 1995). Sintashta-Petrovka communities thus evidence substantial investment of effort in non-subsistence activities, potentially resulting in a subsistence deficit in an economy with a heavy emphasis on herding. Altogether, this makes it plausible to think of the known Sintashta-Petrovka communities as special places where elites for whom military activities were important resided, and where metal production and possibly other crafts were carried out. It remains unclear just how a subsistence economy relying heavily on herding was managed from these substantial sedentary communities. Moving herds around the landscape seasonally is generally thought to be a part of subsistence strategy in Inner Eurasia (Frachetti 2008; Bachura 2013). In this area migration to exploit seasonal pastures is the best strategy for maintaining a regular supply of food for livestock due to shortages of capital or of labor pool to produce, harvest, and store fodder (Dyson-Hudson and Dyson-Hudson 1980:17). The recent stable isotope studies support this notion showing high likelihood that during the Bronze Age livestock was raised locally (Kiseleva et al. 2017).

    The above raises the possibility that the residential remains that have been excavated within the fortifications of Sintashta-Petrovka communities represent only a portion of the population (Hanks and Doonan 2009, Johnson and Hanks 2012). It could be (along with the general lines suggested by D. Zdanovich [1997]) that the archaeological remains of the ordinary people who made up the majority of the population, built the impressive fortifications and stoked the subsistence economy have gone largely undetected. In global comparative perspective, many societies with the features known for Sintashta-Petrovka organization consisted of elite central-place settlements and hinterland populations. In such a scenario, the “missing” portion of the Sintashta population would reside in smaller unfortified settlements scattered around in the vicinity of the fortified ones.

    kamenni-ambar-cultural-layer

    In terms of wealth and productive differentiation, the inside assemblage of Kamennyi Ambar demonstrates a higher degree of richness and diversity in its material assemblage, leading to the conclusion that the outside materials may represent a semi-mobile group of people who used significantly less durable materials and accumulated less possessions. As for the diversity within the inside artifact assemblage, some households at Kamennyi Ambar demonstrate more diverse artifact assemblages than others, as well as bigger sizes, that could be related to differences in productive activities and/or wealth differentiation between families. A focus on specific objects of ceramic production in House 1 suggests some degree of productive specialization, while the elite goods in House 5 clearly point out the presence of elite members of the society.

    There are two possible social scenarios that explain the settlement situation during the Sintashta-Petrovka phase. The first scenario considers all three communities as simultaneous and the second scenario suggests seeing the three sites as the same community that moved around the landscape during the Late Bronze Age in order to keep the pasture grounds from degradation.

    Since no remains of permanent structures were found and any people living outside the walls must have stayed in temporary shelters. If this was the case, then the outside part of the population consisted of a semi-mobile group of people who moved to live near the fortified settlement during the winter. The pattern of animal slaughtering supports this conclusion. Animal teeth found near Kamennyi Ambar and Konoplyanka demonstrate a tendency for animal butchering during the fall, throughout the winter and spring, with less evidence of summer meat consumption. Moreover, since the Bronze Age subsistence strategy relied heavily on pastoralism, herds had to be grazed during the summer and kept safe during the winter. This strongly suggests that the part of the population responsible for management of animals spent their time in the summer pastures with the livestock. During the winter the animals had to be kept in the warm and safe environment of the walled settlements (as suggested by the highest level of phosphorus on the house floors) while the herders stayed in portable shelters in close to the walls.

    (…) the outsiders used a less diverse set of tools, as well as less durable materials (for example, wooden instead of metal) in their everyday life and did not accumulate much in the way of archaeologically visible possessions. On the other hand, a few stone and lithic artifacts demonstrate that craft activities were carried out using cheap and abundant raw materials. The artefact assemblages also point out that the people inside accumulated wealth in the form of material belongings and luxury goods, especially, things like metal artifacts and symbolic or military-related stone artifacts, while people outside did not do that. However, the presence of semi-precious stones could signify some kind of wealth accumulation by the segment of population outside the walls. Since there are limits to our ability to assess social relationships from material remains, it is difficult to say if the people who lived outside the walls were oppressed or less respected. Their possible concentration on herding-related activities and livestock keeping might suggest less prestigious social status. The most prominent members of the society were, nonetheless, buried with the attributes of warriors or craft specialists, not those of shepherds, suggesting that those involved in livestock management had less social prestige.

    Furthermore, Kuzmina (1994:72) cites linguistic studies demonstrating that the Sanskrit word for a permanent village earlier meant a circle of mobile wagon homes, situated together for defensive purposes for an overnight camp (Kuzmina 1994:72).

    The likely population of semi-mobile herders represented some 30%–60% of the entire local community, while the other of 40%–70% were inhabitants of the walled settlement. The almost completely excavated kurgan cemetery of Kamennyi Ambar-5 (only two kurgans remain unstudied) yielded about 100 individuals, or about 2%–5% of the total of 4,896±1,960 individuals in four generations who lived at the nearby settlement for 100 years. In other words, no more than 10% of the population was entitled to be buried under the kurgan mound and this proportion can be taken as an estimate of those with elevated social status. Perhaps, these elites were kin, since analysis of the burial patterns suggests sex/age rather than wealth/prestige differentiation between buried individuals within this elite group (Epimakhov and Berseneva 2011; Ventresca Miller 2013). The remaining non-elite members of the permanently resident community, then, represented some 30%–60% of the complete local community, but did not show evidence of standards of living particularly lower than the elites eventually interred in the kurgan.

    (…) The buried population in the Sintashta Cemetery is about 80 individuals or only about 2%–3% of the total estimated population. However, these few individuals were buried with extremely rich offerings, like complete chariots, decorations made of precious metals or sacrifices of six horses (equal to about 900 kg of meat), etc. With such a low proportion of the population assigned such high prestige, the Sintashta local community can easily be labeled a local chiefdom. In Pitman and Doonan’s view (2018) the social structure of the chifedom consisted of a chief and his kin at the highest level; warriors, religious specialists, and craftsmen in the middle; and the pastoral community at the bottom level.

    kamenni-ambar-excavations

    In the Bronze Age, the people who comprised the majority of the permanent population were involved in craft activities, including extraction of copper ores, metallurgy, bone, leather, and woodwork. The most important and labor-intensive part of the economy, however, was haymaking. The evidence of hay found in the cultural layer near Kamennyi Ambar supports the idea that animals were fed during the winter. Nowadays, hay cutting is typically done in July-August, the period of most intensive grazing for animals. Thus, the part of the collective that remained in the settlement had to provide the labor force for haymaking.

    In the wintertime, the herders returned to the settlements with the herds, and animals were kept inside the walls––a practice which is known archaeologically (Zakh 1995) and ethnographically (Shahack-Gross et al. 2004)––while herders stayed outside in their tents.

    In sum, the Sintashta-Petrovka chiefdoms demonstrate a three-part social order. In Kuzmina’s (1994) view, this is similar to the Varna system of ancient India, that consisted of priests (Sansk. Brahmanis), rulers and warriors (Sansk. Kshatriyas), free producers (Sansk. Vaishyas) and laborers and service providers (Sansk. Shudras). In the Sintashta-Petrovka chiefdom, the elite 2%–5% of the population would have consisted of priests and warriors; 48%–55% would have been dependent producers; and 50%–60% would have been herders of lower social rank.

    sintashta-petrovka-settlements
    The map of the Bronze Age sites in the Karagaily-Ayat Valley Sites of Phase 1: 101 – Konoplyanka; 102 – Zhurumbay; 103 – Kamennyi Ambar; 104 – Kamennyi Ambar-5 Sites of Phase 2: 201 – Konoplyanka 1; 202 – Varshavskoye-1; 203 – Zhurumbay-1; 204 – Varshavskoye-3; 205 – Varshavskoye-5; 206 – Varshavskoye-9; 207 – Kamennyi Ambar-8; 208 – Kamennyi Ambar; 209 – Elizavetpolskoye-3; 210 – Elizavetpolskoye-2; 211 – Karagayli-26; 212 – Elizavetpolskoye-7; 213 – Elizavetpolskoye- 9; 214 – Yuzhno-Stepnoyi (1); 215 – Yuzhno-Stepnoyi (2)

    Conclusions

    In the case of the Sintashta-Petrovka chiefdoms, the questions of why and how exactly social complexity developed through time and why individuals choose to integrate and give up their independence can be answered as some combination of two necessities: to persist as a larger community in the ecological niche of the newly settled region, and to protect herds from theft.

    There is general agreement among researchers that the Sintashta phenomenon had no local roots and originated with a large-scale migration of pastoral communities from Eastern Europe to the marginal area of the Southern Urals. This process forced families to stay together and fueled the necessity in the walled villages for ensuring the reproduction of herds in the extreme climatic conditions of the southern Urals that are colder and dryer than the eastern Black Sea region from which the Sintashta populations are thought to have migrated (Kuzmina 1994, 2007; Anthony 2007; Vinogradov 2011, etc.). At the same time, the herds needed protection from animal and human predators. Probably, the risk of losing animals was a threat to survival that created tensions between neighboring communities, and the Neolithic hunter-gatherers who had populated the Urals before the arrival of Sintashta people could have hunted the domestic animals. Apparently, those who were talented in managing the construction of closely-packed villages surrounded by ditches and walls to protect people and livestock from threats from neighbors, and who otherwise served the community in the newly colonized zone became the most prominent members of society. Theses people formed the core of the Sintashta-Petrovka chiefdom but were not able to accumulate much personal wealth in the form of material possessions. Instead, they acquired high social prestige that could even be transferred to their children (since up to 65% of the buried elite population consists of infants [Razhev and Epimakhov 2005). In this sense, the Sintashta-Petrovka elites were simmilar to their counterparts in the Alto Magdalena of Colombia (Drennan 1995; Gonzalez Fernandez 2007; Drennan and Peterson 2008).

    However, this situation did not last longer than 300 years, since after the initial phase of colonization of the Southern Urals was over, the need for social services provided by an elite disappeared and centralized chiefly communities disintegrated into the smaller unfortified villages of the Srubnaya-Alakul’ period.

    As I have said many times already (see e.g. here) the outsider pastoralists, forming originally the vast majority of the population, were most likely Pre-Proto-Indo-Iranian speakers of haplogroup R1b-Z2103, and their elite groups (whose inheritance system was based on kinship) probably incorporated gradually Uralic-speaking families of haplogroup R1a-Z93, whose relative importance increased gradually, and then eventually expanded massively with the migrations of Andronovo and Srubna, creating a second Y-chromosome bottleneck that favoured again Z93 subclades. The adaptation of Pre-Proto-Indo-Iranian to the Uralic pronunciation, and the adoption of PII vocabulary in neighbouring Proto-Finno-Ugric bear witness to this process.

    Related

    Sintashta diet and economy based on domesticated animal products and wild resources

    indo-iranian-sintashta-uralic-migrations

    New paper (behind paywall) Bronze Age diet and economy: New stable isotope data from the Central Eurasian steppes (2100-1700 BC), by Hanks et al. J. Arch. Sci (2018) 97:14-25.

    Interesting excerpts (emphasis mine):

    Previous research at KA-5 was carried out by A. V. Epimakhov in 1994–1995 and 2002–2003 and resulted in the excavation of three Sintashta culture barrows (kurgans) that produced 35 burial pits and a reported 100 skeletons (Epimakhov, 2002, 2005; Epimakhov et al., 2005; Razhev and Epimakhov, 2004). Seven AMS radiocarbon dates on human remains from the cemetery yielded a date range of 2040–1730 cal. BC (2 sigma), which placed the cemetery within the Sintashta phase of the regional Bronze Age (Hanks et al., 2007). Twelve recently obtained AMS radiocarbon dates, taken from short-lived wood and charcoal species recovered from the Kamennyi Ambar settlement, have provided a date range of 2050–1760 cal. BC (2 sigma). Importantly, these dates confirm the close chronological relationship between the settlement and cemetery for the Middle Bronze Age phase and discount the possibility of a freshwater reservoir effect influencing the earlier dating of the human remains from the Kamennyi Ambar 5 cemetery (Epimakhov and Krause, 2013).

    Sintashta cemeteries frequently yield fewer than six barrow complexes and the number of skeletons recovered represents a fraction of the total population that would have inhabited the settlements (Judd et al., 2018; Johnson and Hanks, 2012). Scholars have suggested that only members of higher status were afforded interment in these cemeteries and that principles of social organization structured placement of individuals within central or peripheral grave pits (Fig. 2) (Koryakova and Epimakhov, 2007: 75–81). In comparison with other Sintashta cemeteries that have been excavated, KA-5 provides one of the largest skeletal inventories currently available for study.

    kamenniy-ambar
    Upper – plan of Kamennyi Ambar settlement and cemetery; Lower – plan views of Kurgan 2 and Kurgan 4 from KA-5 Cemetery (kurgan plans redrawn from Epimakhov, 2005: 10, 79).

    The KA-5 (MBA), Bestamak (MBA) and Lisakovsk (LBA) datasets exhibited a wide range of δ13C and δ15N values for both humans and herbivores (Figs. 5 and 6 & Table 8). This diversity in isotopic signals may be evident for a variety of reasons. For example, the range of values may be associated with a broad spectrum of C3 and C4 plant diversity in the ancient site biome or herbivore grazing patterns that included more diverse environmental niche areas in the microregion around the sampled sites. Herders also may have chosen to graze animals in niche areas due to recognized territorial boundaries between settlements and concomitant patterns of mobility. Importantly, data from Bolshekaragansky represents humans with lower δ15N values that are more closely associated with δ15N values of the sampled domestic herbivores (Fig. 6). When the archaeological evidence from associated settlement sites is considered, Bolshekaragansky, Bestamak, Lisakovsk and KA-5 have been assumed to represent populations that shared similar forms of pastoral subsistence economies with significant dietary reliance upon domesticated herbivore meat and milk. Human diets have δ13C values closely related to those of local herbivores in terms of the slope of the trendline and range of values (Fig. 6). Comparatively, the cemetery of Bolshekaragansky (associated with the Arkaim settlement) reflects individuals with trend lines closer to those of cattle and caprines and may indicate a stronger reliance on subsistence products from these species with less use of wild riverine and terrestrial resources. The site of Čiča is significantly different with elevated human δ15N isotopic values and depleted δ13C values indicative of a subsistence regime more closely associated with the consumption of freshwater resources, such as fish. The stable isotopic data in this instance is strongly supported by zooarchaeological evidence recovered from the Čiča settlement and also is indicative of significant diachronic changes from the LBA phases through the Iron Age (Fig. 6).

    kamenniy-ambar-isotopic-chicha-lisakovsk-bestamark
    Regional analysis and comparison of stable isotope results from humans (adults) and animals recovered from MBA and LBA cemeteries in the Southern Urals (Kamennyi Ambar 5 & Bolshekaragansky) northwestern Kazakhstan (Liskovsk & Bestamak) and southwestern Siberia (Čiča).

    Conclusion

    (…) The isotopic results from KA-5, and recent botanical and archaeological studies from the Kamennyi Ambar settlement, have not produced any evidence for the production or use of domesticated cereals. While this does not definitively answer the question as to whether Sintashta populations engaged in agriculture and/or utilized agricultural products, it does call into serious question the ubiquity of such practices across the region and correlates well with recent archaeological, bioarchaeological, and isotopic studies of human and animal remains from the Southwestern Urals region and Samara Basin (Anthony et al., 2016; Schulting and Richards, 2016). The results substantiate a broader spectrum subsistence diet that in addition to the use of domesticated animal products also incorporated wild flora, wild fauna and fish species. These findings further demonstrate the need to draw on multiple methods and datasets for the reconstruction of late prehistoric subsistence economies in the Eurasian steppes. When possible, this should include datasets from both settlements and associated cemeteries.

    Variability in subsistence practices in the central steppes region has been highlighted by other scholars and appears to be strongly correlated with local environmental conditions and adaptations. More comprehensive isotopic studies of human, animal and fish remains are of fundamental importance to achieve more robust and empirically substantiated reconstructions of local biomes and to aid the refinement of regional and micro-regional economic subsistence models. This will allow for a fuller understanding of key diachronic shifts within dietary trends and highlight regional variation of such practices. Ultimately, this will more effectively index the diverse social and environmental variables that contributed to late prehistoric lifeways and the economic strategies employed by these early steppe communities.

    Social organization of Sintashta-Petrovka

    Interesting to remember now the recent article by Chechushkov et al. (2018) about the social stratificaton in Sintashta-Petrovka, and how it must have caused the long-lasting, peaceful admixture process that led to the known almost full replacement of R1b-L23 (mostly R1b-Z2103) by R1a-Z645 (mostly R1a-Z93) subclades in the North Caspian steppe, coinciding with the formation of the Proto-Indo-Iranian community and language (read my thoughts on this after Damgaard et al. 2018).

    Here is another relevant excerpt from Chechushkov et al. (2018), translated from Russian:

    settlement-kamenniy-ambar
    The map of the settlement of Kamennyi Ambar with excavations, soil cores, and test pits. Legend: a — cuts of the sides of ravines; b — test pits of 2015—2017; c — test pits of 2004; d — soil-science samples with a cultural layer; e — soil-science samples without cultural layer; f — borders of archaeological sites (interpretation of the plan of magnetic anomalies); g — boundaries of excavated structures (1, 2, 4, 5, 7 — Sintashta-Petrovka culture; 3, 6 — Srubnaya-Alakul’ culture).

    The analysis suggests that the Sintashta-Petrovka societies had a certain degree of social stratification, expressed both in selective funeral rituals and in the significant difference in lifestyle between the elite and the immediate producers of the product. The data obtained during the field study suggest that the elite lived within the fortifications, while a part of the population was outside their borders, on seasonal sites, and also in stationary non-fortified settlements. Probably, traces of winter settlements can be found near the walls, while the search for summer ones is a task of a separate study. From our point of view, the elite of the early complex societies of the Bronze Age of the Eurasian steppe originated as a response to environmental challenges that created risks for cattle farming. The need to adapt the team to the harsh and changing climatic conditions created a precedent in which the settled collectives of pastoralists – hunter-gatherers could afford the content and magnificent posthumous celebration of people and their families who were not engaged in the production or extraction of an immediate product. In turn, representatives of this social group directed their efforts to the adoption of socially significant decisions, the organization of collective labor in the construction of settlement-shelters and risked their lives, acting as military leaders and fighters.

    Thus, in Bronze Age steppe societies, the formation, development and decline of social complexity are directly related to the intensity of pastoralism and the development of new territories, where collectives had to survive in part a new ecological niche. At the same time, some members of the collective took upon themselves the organization of the collective’s life, receiving in return a privileged status. As soon as the conditions of the environment and management changed, the need for such functions was virtually eliminated, as a result of which the privileged members of society dissolved into the general mass, having lost their lifetime status and the right to be allocated posthumously.

    Also interesting for the MLBA haplogroup bottleneck in the region is the paper by Judd et al. (2017) about fast life history in Early Indo-Iranian territories.

    On the arrival of haplogroup N1c1-L392

    Regarding the special position of the Chicha-1 samples in the change of diet and economy during the Iron Age, it is by now well known that haplogroup N must have arrived quite late to North-East Europe, and possibly not linked with the expansion of Siberian ancestry – or linked only with some waves of Siberian ancestry in the region, but not all of them. See Lamnidis et al. (2018) for more on this.

    Also, the high prevalence of haplogroup N among Fennic and Siberian (Samoyedic) peoples is not related: while the latter reflects probably the native (Palaeo-Siberian) population that acquired their Uralic branch during the MLBA expansions associated with Corded Ware groups, the former points to the expansion of Fennic peoples into Saamic territory (i.e. after the Fenno-Saamic split) as the most likely period of expansion of N1c1-L392 subclades (see known recent bottlenecks among Finns, and on Proto-Finnic dialectalization).

    Probably related to these late incomers are the ancient DNA samples from the Sargat culture during the Iron Age, which show the arrival of N subclades in the region, replacing most – but not all – R1a lineages (see Pilipenko et al. (2017)). Regarding the site of Chicha-1, the following are relevant excerpts about the cultural situation that could have allowed for such stepped, diachronic admixture events in Northern Eurasia, from the paper Stages in the settlement history of Chicha-1: The Results of ceramic analysis, by Molodin et al. (2008):

    The stratigraphic data allows us to make the following inference: originally, the settlement was inhabited by people bearing the Late Irmen culture. Later, the people of the Baraba trend of the Suzgun culture arrived at the site (Molodin, Chemyakina, 1984: 40–62). The Baraba-Suzgun pottery demonstrates features similar to what has been reported from the sites of the transitional Bronze to Iron Age culture in the pre-taiga and taiga zones in the Irtysh basin (Potemkina, Korochkova, Stefanov, 1995; Polevodov, 2003). The major morphological types are slightly and well-profiled pots with a short throat. (…)

    chicha-irmen-tagar-baraba-forest-siberian
    Map showing the location of Chicha-1.

    During the following stage of development of the site, the Chicha population increased with people who practiced cultures others than those noted in earlier collections. The ceramic materials from layer 5 provide data on possible relationships. In addition to migrants from northwestern regions practicing the Suzgun culture, there were people bearing the Krasnoozerka culture. Available data also suggests that people from the northern taiga region with the Atlym culture visited the site.

    However, people from the west and southwest represent the greatest migration to the region under study. In all likelihood they moved from the northern forest-steppe zone of modern Kazakhstan and practiced the Berlik culture. The spatial distribution analysis of the Chicha-1 site suggests that the Berlik population was rather large. The Berlik people formed a single settlement with the indigenous Late Irmen people and apparently waged certain common economic activities, but preserved their own ethnic and cultural specificity (Molodin, Parzinger, 2006: 49–55). Judging by the data on the chronological sequence of deposited artifacts, migration took place roughly synchronously, hence Chicha-1 became a real cultural and economic center.

    (…) In sum, the noted distribution of ceramics over the culture-bearing horizons suggests that beginning with layer 5, traditions of ceramic manufacture described above were practiced, hence the relevant population inhabited the site. Apparently, there were two predominant traditions: the local Late Irmen cultural tradition and the Berlik tradition, which was brought by the immigrants. The Late Irmen people mostly populated the citadel, while the Berlik immigrants inhabited the areas to the east and the north of the citadel.

    The stratigraphic data also suggest that the Early Sargat ceramics emerged at the site likely as a part of the Late Irmen tradition (…) Early Sargat ceramics is apparently linked with the Late Irmen tradition. Artifacts associated with the Sargat culture proper have been found in several areas of Chicha-1 (e.g., in excavation area 16). However, the Sargat people appeared at the site after it had been abandoned by its previous inhabitants, and had eventually become completely desolated. This happened no earlier than the 6th cent. BC, possibly in the 5th cent. BC (in fact, the radiocarbon dates for that horizon are close to the turn of the Christian era).

    Related

    Genetic history of admixture across inner Eurasia; Botai shows R1b-M73

    y-haplogroup-r1b-p343

    Open access Characterizing the genetic history of admixture across inner Eurasia, by Jeong et al. (2018).

    Abstract (emphasis mine):

    The indigenous populations of inner Eurasia, a huge geographic region covering the central Eurasian steppe and the northern Eurasian taiga and tundra, harbor tremendous diversity in their genes, cultures and languages. In this study, we report novel genome-wide data for 763 individuals from Armenia, Georgia, Kazakhstan, Moldova, Mongolia, Russia, Tajikistan, Ukraine, and Uzbekistan. We furthermore report genome-wide data of two Eneolithic individuals (~5,400 years before present) associated with the Botai culture in northern Kazakhstan. We find that inner Eurasian populations are structured into three distinct admixture clines stretching between various western and eastern Eurasian ancestries. This genetic separation is well mirrored by geography. The ancient Botai genomes suggest yet another layer of admixture in inner Eurasia that involves Mesolithic hunter-gatherers in Europe, the Upper Paleolithic southern Siberians and East Asians. Admixture modeling of ancient and modern populations suggests an overwriting of this ancient structure in the Altai-Sayan region by migrations of western steppe herders, but partial retaining of this ancient North Eurasian-related cline further to the North. Finally, the genetic structure of Caucasus populations highlights a role of the Caucasus Mountains as a barrier to gene flow and suggests a post-Neolithic gene flow into North Caucasus populations from the steppe.

    Interesting excerpts:

    On North Eurasians

    In a PCA of Eurasian individuals, we find that PC1 separates eastern and western Eurasian populations, PC2 splits eastern Eurasians along a north-south cline, and PC3 captures variation in western Eurasians with Caucasus and northeastern European populations at opposite ends (Figure 2A and Figures S1-S2). Inner Eurasians are scattered across PC1 in between, largely reflecting their geographic locations. Strikingly, inner Eurasian populations seem to be structured into three distinct west-east genetic clines running between different western and eastern Eurasian groups, instead of being evenly spaced in PC space. Individuals from northern Eurasia, speaking Uralic or Yeniseian languages, form a cline connecting northeast Europeans and the Uralic (Samoyedic) speaking Nganasans from northern Siberia (“forest-tundra” cline). Individuals from the Eurasian steppe, mostly speaking Turkic and Mongolic languages, are scattered along two clines below the forest-tundra cline. Both clines run into Turkic- and Mongolic-speaking populations in southern Siberia and Mongolia, and further into Tungusic-speaking populations in Manchuria and the Russian Far East in the East; however, they diverge in the west, oneheading to the Caucasus and the other heading to populations of the Volga-308 Ural area (the “southern steppe” and “steppe-forest” clines, respectively; Figure 2 and Figure S2).
    (…)
    The forest-tundra cline populations derive most of their eastern Eurasian ancestry from a component most enriched in Nganasans, while those on the steppe-forest and southern steppe clines have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh. The southern steppe cline groups are distinct from the others in their western Eurasian ancestry profile, in the sense that they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers (“CHG”) and Neolithic Iranians (“Iran_N”) and frequently harbor another component enriched in South Asians (Figure S4).

    north-eurasian-uralic
    qpAdm-based admixture models for the forest-tundra cline populations. For populations to the east of the Urals (Enets, Selkups, Kets, and Mansi), EHG+Yamnaya+Nganasan provides a good fit, except for Mansi, for which adding WHG significantly increases the model fit. For the rest of the groups, WHG+LBK_EN+Yamnaya+Nganasan in general provides a good fit. 5 cM jackknifing standard errors are marked by the horizontal bar.

    For the forest-tundra cline populations, for which currently no relevant Holocene ancient genomes are available, we took a more generalized approach of using proxies for contemporary Europeans: WHG, WSH (represented by “Yamnaya_Samara”), and early Neolithic European farmers (EEF; represented by “LBK_EN”; Table S2). Adding Nganasans as the fourth reference, we find that most Uralic-speaking populations in Europe (i.e. west of the Urals) and Russians are well modeled by this four-way admixture model (χ 2 p ≥ 0.05 for all but three groups; Figure 5 and Table S8). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in model fit (4.7% to 29.1% contribution; χ 2 p ≤ 1.12×10-8; Table S8). The ratio of contributions from three European references varies from group to group, probably reflecting genetic exchange with neighboring non-Uralic groups. For example, Saami from northern Fennoscandia contain a higher WHG and lower WSH contribution (16.1% and 41.3%, respectively) than Udmurts or Besermyans from the Volga river region do (4.9-6.6% and 50.7-53.2%, respectively), while the three groups have similar amounts of Nganasan-related ancestry (25.5-29.1%).

    The Caucasus Mountains form a barrier to gene flow

    By applying EEMS to the Caucasus region, we identify a strong barrier to gene flow separating North and South Caucasus populations. This genetic barrier coincides with the Greater Caucasus mountain ridge even to small scale: a weaker barrier in the middle, overlapping with Ossetia, matches well with the region where the ridge also becomes narrow. We also observe weak barriers running in the north-south direction that separate northeastern populations from northwestern ones. Together with PCA, EEMS results suggest that the Caucasus Mountains have posed a strong barrier to human migration.

    caucasus-genetic-barrier
    The Greater Caucasus mountain ridge as a barrier to 856 genetic exchange. Barriers (brown) and conduits (green) of gene flow around the Caucasus region are estimated by the EEMS program. Red diamonds show the location of vertices to which groups are assigned. A strong barrier to gene flow overlaps with the Greater Caucasus mountain ridge reflecting the genetic differentiation between populations of the north and south of the Caucasus. The barrier becomes considerably weaker in the middle where present-day Ossetians live.

    On the Botai individuals

    The Y-chromosome of the male Botai individual (TU45) belongs to the haplogroup R1b (Table 411 S6). However, it falls into neither a predominant European branch R1b-L5165 nor into a R1b-GG400 branch found in Yamnaya individuals. Thus, phylogenetically this Botai individual should belong to the R1b-M73 branch which is frequent in the Eurasian steppe (Figure S9). This branch was also found in Mesolithic samples from Latvia as well as in numerous modern southern Siberian and Central Asian groups.

    The Botai genomes provide a critical snapshot of the genetic profile of pre-Bronze Age steppe populations. Our admixture modeling positions Botai primarily on an ancient genetic cline of the pre-Neolithic western Eurasian hunter-gatherers: stretching from the post-Ice Age western European hunter-gatherers (e.g. WHG) to EHG in Karelia and Samara to the Upper Paleolithic southern Siberians (e.g. AG3). Botai’s position on this cline, between EHG and AG3, fits well with their geographic location and suggests that ANE-related ancestry in the East did have a lingering genetic impact on Holocene Siberian and Central Asian populations at least till the time of Botai.
    (…)
    The most recent clear connection with the Botai ancestry can be found in the Middle Bronze Age Okunevo individuals (Figure S6C). In contrast, additional EHG-related ancestry is required to explain the forest-tundra populations to the east of the Urals (Figure 5 and Table S8). Their multi-way mixture model may in fact portrait a prehistoric two-way mixture of a WSH population and a hypothetical eastern Eurasian one that has an ANE-related contribution higher than that in Nganasans. Botai and Okunevo individuals prove the existence of such ANE ancestry-rich populations. Pre-Bronze Age genomes from Siberia will be critical for testing this hypothesis.

    botai-pca
    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 north-south 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.

    So, to sum up:

    • Northern Eurasia forms a Uralic – Yeniseian cline from east to west, with contribution from Steppe, WHG, and Siberian ancestry. Siberian ancestry is represented by Palaeo-Siberian Nganasans, who adopted Samoyedic quite late. It was already known that the different waves of Siberian ancestry are too late and do not represent the spread of Uralic languages, so that leaves us with Steppe and WHG.
    • The Caucasus Mountains were a long-lasting prehistoric barrier to gene flow (as recently shown in Y-DNA, too).
    • The Botai sample (ca. 3632-3100 BC) represents thus the furthest east that R1b-P297 subclades had expanded (we did know that, and that they didn’t have close genetic links with Khvalynsk, so the haplogroup spread there probably much earlier). It expanded R1b-M269’s sister clade R1b-M73 (also found in the Baltic region), and the Botai are on the ‘eastern’ end of an ancient genetic cline stretching from WHG to EHG to Afontova Gora.

    EDIT (23 MAY 2018) Both samples share mtDNA, and the male one shares Y-DNA, with those reported in Damgaard et al. (Nature 2018); although dates are slightly different (3371-3354 calBC for BOT 14), it is within the range given for this one; for the female, the dates are similar (3521-3377 calBC for BOT2016, 3517-3367 cal. BCE for this one). The lack of data on their origin may point to the fact that we only have different bone samples from the same two Botai individuals. So probably still 50% R1b-M73 (with the other 50% being N2* from BOT15)…

    It seems therefore not only that R1b-M269 is bound to split from the parent haplogroup in or around the steppe or forest-steppe: the Mesolithic spread of haplogroup R1b in North Eurasia is wider and its relevance thus greater than previously thought.

    We may need to rethink the role of haplogroup R1a in spreading EHG and Indo-Uralic from east to west…

    Featured image, from the supplementary materials: Frequency distribution map of the Y-chromosomal haplogroup R1b-P343(xM269) identified in the Eneolithic Botai individual. All modern Eurasian samples with this haplogroup tested to date for the downstream markers fall into R1b-M73 branch, suggesting Botai sample be one of its earliest representatives.

    Related:

    North-Eastern Europe in the Stone Age – bridging the gap between the East and the West

    chalcolithic_early_corded_ware

    Interesting PhD thesis The Stone Age of north-eastern Europe 5500–1800 calBC : bridging the gap between the East and the West by Kerkko Nordqvist (2018).

    Some interesting excerpts:

    On the Corded Ware and related cultures

    The arrival of Corded Ware is without a doubt the clearest example of migration recognized in Finnish Stone Age archaeology. Its appearance has been understood to result from the movement of a new population from the southern or southeastern Baltic Sea area to the southern and western coasts of Finland (Europaeus 1922: 137; Luho 1948: 57; Edgren 1970: 62; Matiskainen 1994: 14) (Fig. 36). Native inhabitants of the coastal region — presented as representatives of Comb Ware population — have rarely been given any larger role in this development (Luoto 1986: 19; Asplund 1995: 74; see also Article III), and their fate has usually been described as displacement, assimilation or some kind of co-existence (Äyräpää 1952a: 24–25; Edgren 1997: 169–171; Carpelan 1999: 263–264; Núñez 2004: 362).

    Another important part of the Finnish narrative is the distance that the Corded Ware population assumedly kept, not only from the (Pöljä Ware-producing) hunter-gatherers inhabiting the Finnish inland, but also from the other Corded Ware groups in the northern Baltic Sea area (Edgren 1970: 61; Äyräpää 1973: 199, 207; Carpelan 1999: 266).56 The image of Finnish Corded Ware is static: it is seen to exist in the area of present-day Finland facing little change for centuries (Edgren 1992: 96; see also Luoto 1986: 17; Matiskainen 1994). However, the archaic nature assigned to Corded Ware derives greatly from the out-dated idea of pan-European A-horizon, unrealistic dating given for the phenomenon, as well as an overly narrow view of cultural dynamics concerning what can be accepted as Corded Ware (cf. Furholt 2014; Article III).

    The only larger change has been connected to the so-called 2nd wave of Corded Ware, supposedly reaching Finnish coasts from Estonia towards the end of Corded Ware’s existence. However, this event has never really been substantiated, and in Finnish assemblages it seems to materialize only through the so-called sharp-butted axes (see Soikkeli 1912; Äyräpää 1952b: 89–90) (Fig. 37) — pottery related to the 2nd wave has never been presented, although its influences are recognized in later pottery types (Carpelan 1979: 15; Carpelan et al. 2008: 206; see also Lavento 2001: 24–25). Along the northern limit of Corded Ware, in the so-called Middle Zone, the 2nd wave assumedly contributed to the creation of hybrid pottery, which in the earlier research has been vaguely called Middle or Intermediate Zone Ceramics (Carpelan 1979: 15; 2004b: 52). More recently it has been proposed that such mixing of influences and hybridization would have started immediately or soon after the arrival of Corded Ware at least on the south-eastern coast and the Karelian Isthmus, and influences would have been transmitted towards the inland and the middle-zone, too (see Mökkönen 2011: 62–63; Article III; see also Carpelan 1999: 262).

    asbestos-tempered-wares
    Distribution of the so-called Middle and Late Neolithic asbestos- and organictempered wares. Organic admixtures were commonly used in wide regions to the east and south-east of the research area during this time as well, but asbestos-tempered pottery has only occasionally been reported from areas further east of Lake Onega, the Vologda and Arkhangelsk Oblasts (see Ошибкина 1978; Козырева 1983; Жульников 2007). Illustration: K. Nordqvist.

    On assumed early Corded Ware materials in North-East Europe

    The emergence of Corded Ware was previously dated in Finland as early as 3200 calBC (Edgren 1992: 92; Matiskainen 1994: 14; Carpelan 2004b: 48–49). The age was based on a few conventional dates from mixed contexts, and has been lately readjusted to around 2900–2800 calBC (Mökkönen 2011: 17–18; Article III). This has not only moved the dating closer to the initial dates given to Corded Ware in Europe (Włodarczak 2009), but also changed the cultural context into which Corded Ware may have arrived in north-eastern Europe. The old dating permitted the assumption of a temporal overlap with Typical/Late Comb Ware (see Edgren 1970: 59–60; see also Carpelan 1999: 262) — with regards to the new dating, even if the Comb Ware tradition continued in some form to the 3rd millennium calBC, it is really not known how the coastal societies transformed and what they looked like during this time. All in all, only a few sites have been securely dated to the 3200–2800 calBC period, which makes estimating all the mechanisms through which Corded Ware was established quite complicated.

    Most Corded Ware materials derive from mixed, multi-period settlement contexts, which explain the generally limited knowledge about Corded Ware assemblages. Pottery is the most commonly identified element, although its study has been heavily concentrated on beakers and beaker-like cups: apart from so-called short-wave moulded vessels, household pottery is not much recognized (Edgren 1970: 25–26; see Nordqvist & Häkälä 2014: 18–19). Furthermore, apart from individual remarks, organic tempers have been excluded from the Corded Ware technological repertoire in Finland (Edgren 1970: 33; Korkeakoski- Väisänen 1993: 15) — as shown in Article III, organic-tempered Corded Ware is present at least on the Finnish southern coast and the Karelian Isthmus, and has been reported from Southern Ostrobothnia as well. Organic-tempered pottery found in southern Finland is similar to the so-called Estonian (or Late) Corded Ware, which is thought to be the result of local development (Янитс 1959: 166; Kriiska 2000: 75; see also Kholkina 2017: 155) (Fig. 38). Even preliminary mapping of such pottery (Finnish data is still based on non-systematic survey) shows that an interaction sphere existed in the eastern Gulf of Finland area, reaching from Estonia to the areas of present-day Finland and the Karelian Isthmus in Russia (Article III) (Fig. 36). Sharp-butted axes fit well into this context: rather than being an indication of some ambivalent and unidirectional 2nd wave of influence, they provide better evidence of more continuous contacts across the sea.

    The origins and spread of Corded Ware have become highly topical in the last few years with the development of analytical techniques such as genetic and isotopic research (Allentoft et al. 2015; Haak et al. 2015; Sjögren et al. 2016; Kristiansen et al. 2017). Generally, archaeogenetic studies have evidenced large population replacements in Europe, and seem to provide solid support for migration — still, numerous problems related to representativity and interpretation of the data remain to be solved (see Vander Linden 2016; Heyd 2017; Ion 2017). No material is available for such studies from the research area, as no bones have been preserved in the excavated burials. The closest analysed and published individuals from Estonia, Latvia and Lithuania (Allentoft et al. 2015; Jones et al. 2017; Mittnik et al. 2018; Saag et al. 2017) show that the development of Corded Ware in the Baltic States clearly involved newcomers. At the same time, archaeological materials from north-eastern Europe also indicate the input and presence of indigenous people — settling this discrepancy between different source materials is an important task for future research.

    On modern political borders and preconceptions defining archaeological cultures

    The results presented above illustrate how the modern political borders may appear to be present in the past. For Finnish prehistory they mean also that instead of one ‘Finnish’ group, there are several Corded Ware populations operational within the present-day state. Recent geochemical analyses of clay pastes and grog tempers of Corded Ware have pointed towards the existence of different pottery recipes in different parts of Finland, as well as towards connections and movement across the northern Baltic Sea area (Holmqvist et al. 2018).

    corded-ware-finland-estonia
    The area of Battle Axe/Corded Ware cultures (shaded), with the traditional limit of Finnish Corded Ware shown as a solid line. The distribution of organic-tempered (Estonian) Corded Ware is marked with a dashed line, while the so-called Middle Zone is indicated roughly by hatching. Illustration: K. Nordqvist.

    In the present-day Russian territory, a Corded Ware presence has been recognized on the Karelian Isthmus (Крайнов 1987b: 61, Карта 6). In Finnish archaeology this area has been mostly considered a periphery of Finnish Corded Ware (Äyräpää 1952a: 22–23; 1973: 207; Meinander 1954a: 151–152; Huurre 2003: 236; Carpelan et al. 2008: 206), but the identification of organic-tempered Cored Ware in areas north of the Gulf (as well as mapping of all stray finds; Nordqvist & Häkälä 2014; Article III) shows that a Corded Ware presence on the Isthmus was stronger than thought. In other words, the normative perception of culture and strong presuppositions of what Corded Ware should be has led to the exclusion of part of the material culturethe Karelian Isthmus was not just a subsidiary area of Finnish Corded Ware, but a region with its own character and tradition.

    No Corded Ware pottery finds have been reported in the areas north of Lake Ladoga. The solitary Corded Ware influences noted in simultaneous Karelian pottery have been connected with the central Russian Fatyanovo culture (Жульников 1999: 53–54; 2008: 419). Because the Fatyanovo territory extends close to the research area in the east and south-east (see Крайнов 1987b: 61, Карта 6; see also Жульников 2008: 417, Рис. 3), it is not surprising that recent studies have revealed evidence of connections between the eastern Gulf of Finland and central Russian battle axe cultures (Kriiska et al. 2015: 47; Крийска et al. 2015: 201; Article III; see also Kholkina 2017: 154–155).

    People prefer ethnolinguistic identification “large and early”

    The East has been mostly contextualized through the cultural entity perceived to exist between the Baltic Sea and the Urals, i.e. the ‘Comb Ware cultural sphere’. Its binding elements have been kinship- and exchange-based connections, which would have transmitted influences over this vast area. Karelia, the closest-lying region of the East, has been occasionally mentioned as an important area of influence (Tallgren 1938b; Äyräpää 1944), but generally it has not been given any prominent position in the narratives. In fact, Karelia and much of the Comb Ware sphere gained their insignia and paraphernalia already during the first half of the 20th century, and only fairly petrified stereotypes have been presented in the subsequent literature.

    On the arrival of the Metal Ages, influence from Pre-Proto-Germanic, and the reasons for the genetic bottleneck

    Periodization-wise the question is straightforward: the Neolithic ends with the onset of the Early Metal Period, the Eneolithic (in Russia) and the Bronze Age (in Finland).

    A much larger problem affecting the study of transition has been the general decrease and even lack of archaeological material pertaining to this time. This situation prevails in large areas from the later 3rd millennium calBC onwards and is accentuated during the 2nd millennium calBC. The disappearance of archaeological evidence has been explained by decreasing population numbers which would have been caused by the deteriorating climate (Lavento 2015: 125; see also Sundell 2014). Nevertheless, in the territory of present-day Finland the abundant number of burial cairns (see Meinander 1954b: 89–120; Saipio 2011) as well as pollen analyses showing anthropogenic activities dating to this time (see Alenius et al. 2009; Augustson et al. 2013) indicate that no complete depopulation took place. Therefore, in addition to sparse habitation, the change must be explained also through changing ways of living and material cultures, which make the material remains more difficult to identify archaeologically (also Lavento 2015: 125, 132).

    The changes taking place during this time seem to be connected to external influences. On the coast, the Kiukainen culture is thought to have transformed under Scandinavian influences into the so-called Western Bronze Age, exhibiting changes in their settlements, material culture, means of subsistence and their world view (Meinander 1954b: 196–197; Lavento 2015: 198–199). Development further east, in the areas previously occupied by populations producing asbestos and organic-tempered wares, is characterized by the appearance of so-called Textile Ware, apparently introduced there by a new population originating from the south-east and ultimately from the Volga region (Meinander 1954b; Гурина 1961; Косменко 1992; Lavento 2001). Even though it is not clear what the relationships between the carriers of this new tradition and the local populations were (did the latter perish, assimilate, or coexist?), it is evident that changes took place in all fields of life — and the traditional image of an archaic, static inland is not considered correct anymore (see Saipio 2008; Lavento 2015). However, this does not imply one synchronous or abrupt change or a complete turnover but, for example, traditional forms of subsistence held their ground alongside (slash and burn) agriculture for centuries, even millennia to come.

    All in all, a complex account of events in North-East Europe that will define the ethnolinguistic identification of Corded Ware migrants.

    Related: