An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.
Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).
1. Samara to Early Khvalynsk
The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).
Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.
This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:
NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.
2. Early Khvalynsk expansion
We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).
We also have indirect data. First, there is the PCA with outliers:
Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).
Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).
3. Proto-Corded Ware expansion
It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.
Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).
Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.
NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.
The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.
4. Repin / Early Yamna expansion
We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.
Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:
This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:
We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:
The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.
NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.
A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.
NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.
Interesting excerpts (emphasis mine, some links to images and tables deleted for clarity):
Late Bronze Age (LBA) Srubnaya-Alakulskaya individuals carried mtDNA haplogroups associated with Europeans or West Eurasians (17) including H, J1, K1, T2, U2, U4, and U5 (table S3). In contrast, the Iron Age nomads (Cimmerians, Scythians, and Sarmatians) additionally carried mtDNA haplogroups associated with Central Asia and the Far East (A, C, D, and M). The absence of East Asian mitochondrial lineages in the more eastern and older Srubnaya-Alakulskaya population suggests that the appearance of East Asian haplogroups in the steppe populations might be associated with the Iron Age nomads, starting with the Cimmerians.
#UPDATE (5 OCT 2018): Some Y-SNP calls have been published in a Molgen thread, with:
Srubna samples have possibly two R1a-Z280, three R1a-Z93.
Cimmerians may not have R1b: cim357 is reported as R1a.
Some Scythians have low coverage to the point where it is difficult to assign even a reliable haplogroup (they report hg I2 for scy301, or E for scy197, probably based on some shared SNPs?), but those which can be reliably assigned seem R1b-Z2103 [hence probably the use of question marks and asterisks in the table, and the assumption of the paper that all Scythians are R1b-L23]:
The most recent subclade is found in scy305: R1b-Z2103>Z2106 (Z2106+, Y12538/Z8131+)
scy304: R1b-Z2103 (M12149/Y4371/Z8128+).
scy009: R1b-P312>U152>L2 (P312+, U152?, L2+)?
Sarmatians are apparently all R1a-Z93 (including tem002 and tem003);
Srubnaya-Alakulskaya individuals exhibited genetic affinity to northern and northeastern present-day Europeans, and these results were also consistent with outgroup f3 statistics.
The Cimmerian individuals, representing the time period of transition from Bronze to Iron Age, were not homogeneous regarding their genetic similarities to present-day populations according to the PCA. F3 statistics confirmed the heterogeneity of these individuals in comparison with present-day populations
The Scythians reported in this study, from the core Scythian territory in the North Pontic steppe, showed high intragroup diversity. In the PCA, they are positioned as four visually distinct groups compared to the gradient of present-day populations:
A group of three individuals (scy009, scy010, and scy303) showed genetic affinity to north European populations (…).
A group of four individuals (scy192, scy197, scy300, and scy305) showed genetic similarities to southern European populations (…).
A group of three individuals (scy006, scy011, and scy193) located between the genetic variation of Mordovians and populations of the North Caucasus (…). In addition, one Srubnaya-Alakulskaya individual (kzb004), the most recent Cimmerian (cim357), and all Sarmatians fell within this cluster. In contrast to the Scythians, and despite being from opposite ends of the Pontic-Caspian steppe, the five Sarmatians grouped close together in this cluster.
A group of three Scythians (scy301, scy304, and scy311) formed a discrete group between the SC and SE and had genetic affinities to present-day Bulgarian, Greek, Croatian, and Turkish populations (…).
Finally, one individual from a Scythian cultural context (scy332) is positioned outside of the modern West Eurasian genetic variation (Fig. 1C) but shared genetic drift with East Asian populations.
The presence of an SA component (as well as finding of metals imported from Tien Shan Mountains in Muradym 8) could therefore reflect a connection to the complex networks of the nomadic transmigration patterns characteristic of seasonal steppe population movements. These movements, although dictated by the needs of the nomads and their animals, shaped the economic and social networks linking the outskirts of the steppe and facilitated the flow of goods between settled, semi-nomadic, and nomadic peoples. In contrast, all Cimmerians carried the Siberian genetic component. Both the PCA and f4 statistics supported their closer affinities to the Bronze Age western Siberian populations (including Karasuk) than to Srubnaya. It is noteworthy that the oldest of the Cimmerians studied here (cim357) carried almost equal proportions of Asian and West Eurasian components, resembling the Pazyryks, Aldy-Bel, and Iron Age individuals from Russia and Kazakhstan (12). The second oldest Cimmerian (cim358) was also the only one with both uniparental markers pointing toward East Asia. The Q1* Y chromosome sublineage of Q-M242 is widespread among Asians and Native Americans and is thought to have originated in the Altai Mountains (24)
In contrast to the eastern steppe Scythians (Pazyryks and Aldy-Bel) that were closely related to Yamnaya, the western North Pontic Scythians were instead more closely related to individuals from Afanasievo and Andronovo groups. Some of the Scythians of the western Pontic-Caspian steppe lacked the SA and the East Eurasian components altogether and instead were more similar to a Montenegro Iron Age individual (3), possibly indicating assimilation of the earlier local groups by the Scythians.
Toward the end of the Scythian period (fourth century CE), a possible direct influx from the southern Ural steppe zone took place, as indicated by scy332. However, it is possible that this individual might have originated in a different nomadic group despite being found in a Scythian cultural context.
I am surprised to find this new R1b-L23-based bottleneck in Eastern Iranian expansions so late, but admittedly – based on data from later times in the Pontic-Caspian steppe near the Caucasus – it was always a possibility. The fact that pockets of R1b-L23 lineages remained somehow ‘hidden’ in early Indo-Iranian communities was clear already since Narasimhan et al. (2018), as I predicted could happen, and is compatible with the limited archaeological data on Sintashta-Potapovka populations outside fortified settlements. I already said that Corded Ware was out of Indo-European migrations then, this further supports it.
Even with all these data coming just from a north-west Pontic steppe region (west of the Dnieper), these ‘Cimmerians’ – or rather the ‘Proto-Scythian’ nomadic cultures appearing before ca. 800 BC in the Pontic-Caspian steppes – are shown to be probably formed by diverse peoples from Central Asia who brought about the first waves of Siberian ancestry (and Asian lineages) seen in the western steppes. You can read about a Cimmerian-related culture, Anonino, key for the evolution of Finno-Permic peoples.
Also interesting about the Y-DNA bottleneck seen here is the rejection of the supposed continuous western expansions of R1a-Z645 subclades with steppe tribes since the Bronze Age, and thus a clearest link of the Hungarian Árpád dynasty (of R1a-Z2123 lineage) to either the early Srubna-related expansions or – much more likely – to the actual expansions of Hungarian tribes near the Urals in historic times.
NOTE. I will add the information of this paper to the upcoming post on Ugric and Samoyedic expansions, and the late introduction of Siberian ancestry to these peoples.
A few interesting lessons to be learned:
Remember the fantasy story about that supposed steppe nomadic pastoralist society sharing different Y-DNA lineages? You know, that Yamna culture expanding with R1b from Khvalynsk-Repin into the whole Pontic-Caspian steppes and beyond, developing R1b-dominated Afanasevo, Bell Beaker, and Poltavka, but suddenly appearing (in the middle of those expansions through the steppes) as a different culture, Corded Ware, to the north (in the east-central European forest zone) and dominated by R1a? Well, it hasn’t happened with any other steppe migration, so…maybe Proto-Indo-Europeans were that kind of especially friendly language-teaching neighbours?
Remember that ‘pure-R1a’ Indo-Slavonic society emerged from Sintashta ca. 2100 BC? (or even Graeco-Aryan??) Hmmmm… Another good fantasy story that didn’t happen; just like a central-east European Bronze Age Balto-Slavic R1a continuitydidn’t happen, either. So, given that cultures from around Estonia are those showing the closest thing to R1a continuity in Europe until the Iron Age, I assume we have to get ready for the Gulf of Finland Balto-Slavic soon.
Remember that ‘pure-R1a’ expansion of Indo-Europeans based on the Tarim Basin samples? This paper means ipso facto an end to the Tarim Basin – Tocharian artificial controversy. The Pre-Tocharian expansion is represented by Afanasevo, and whether or not (Andronovo-related) groups of R1a-Z645 lineages replaced part or eventually all of its population before, during, or after the Tocharian expansion into the Tarim Basin, this does not change the origin of the language split and expansion from Yamna to Central Asia; just like this paper does not change the fact that these steppe groups were Proto-Iranian (Srubna) and Eastern Iranian (Scythian) speakers, regardless of their dominant haplogroup.
Do you smell that fresher air? It’s the Central and East European post-Communist populist and ethnonationalist bullshit (viz. pure blondR1a-based Pan-Nordicism / pro-Russian Pan-Slavism / Pan-Eurasianism, as well as Pan-Turanism and similar crap from the 19th century) going down the toilet with each new paper.
#EDIT (5 OCT 2018): It seems I was too quick to rant about the consequences of the paper without taking into account the complexity of the data presented. Not the first time this impulsivity happens, I guess it depends on my mood and on the time I have to write a post on the specific work day…
While the data on Srubna, Cimmerians, and Sarmatians shows clearer Y-DNA bottlenecks (of R1a-Z645 subclades) with the new data, the Scythian samples remain controversial, because of the many doubts about the haplogroups (although the most certain cases are R1b-Z2103), their actual date, and cultural attribution. However, I doubt they belong to other peoples, given the expansionist trends of steppe nomads before, during, and after Scythians (as shown in statistical analyses), so most likely they are Scythian or ‘Para-Scythian’ nomadic groups that probably came from the east, whether or not they incorporated Balkan populations. This is further supported by the remaining R1b-P312 and R1b-Z2103 populations in and around the modern Eurasian steppe region.
You can find an interesting and detailed take on the data published (in Russian) at Vol-Vlad’s LiveJournal (you can read an automatic translation from Google). I think that post is maybe too detailed in debunking all information associated to the supposed Scythians – to the point where just a single sample seems to be an actual Scythian (?!) -, but is nevertheless interesting to read the potential pitfalls of the study.
In recent decades, evidence has accumulated for comparable enclosures of later dates, including the Early Bronze Age Únětice Culture between 2200 and 1600 BC, and thus into the chronological and cultural context of the Nebra sky disc. Based on the analysis of one of these enclosure sites, recently excavated at Pömmelte on the flood plain of the Elbe River near Magdeburg, Saxony-Anhalt, and dating to the late third millennium BC
The main occupation began at 2321–2211 cal BC, with the stratigraphically earliest features containing exclusively Bell Beaker finds. Bell Beaker ceramics continue after 2204–2154 cal BC (boundary occupation I/II), although they were probably undecorated, but are now complemented by Únětice Culture (and other Early Bronze Age) types. At this time, with features common to both cultures predominate. Only contexts dating to the late main occupation phase (late phase II) and thereafter contained exclusively Únětice Culture finds. Evidently, the bearers of the Bell Beaker Culture were the original builders of the enclosure. During a second phase of use, Final Neolithic and Early Bronze Age cultures coexisted and intermingled. The material remains, however, should not be taken as evidence for successive groups of differing archaeological cultures, but as witnesses to a cultural transition from the Bell Beaker Culture to the Únětice Culture (Spatzier 2015). The main occupation ended 2086–2021 cal BC with the deconstruction of the enclosure; Bell Beaker finds are now absent. Finally, a few features (among them one shaft) and radiocarbon dates attest the sporadic re-use of the site in a phase of abandonment/re-use that ended 1636– 1488 cal BC.
How the above-ground structures possibly influenced perception may reveal another layer of meaning that highlights social functions related to ritual. While zone I was disconnected from the surroundings by a ‘semi-translucent’ post-built border, zones II/III were separated from the outside world by a wooden wall (i.e. the palisade), and zone III probably separated individuals from the crowd gathered in zone II. Accessing the interior or centre therefore meant passing through transitional zones, to first be secluded and then segregated. Exiting the structure meant re-integration and re-connection. The experience possibly induced when entering and leaving the monument reflects the three stages of ‘rites of passage’ described by van Gennep (1909): separation, liminality and incorporation. The enclosure’s outer zone(s) represents the pre- and post-liminal phase; the central area, the liminal phase. Seclusion and liminality in the interior promoted a sense of togetherness, which can be linked to Turner’s “communitas” (1969: 132–33). We might therefore see monuments such as the Pömmelte enclosure as important communal structures for social regulation and the formation of identity.
(…) The long-term stability of these connotations must be emphasised. As with the tradition of making depositions, these meanings were valid from the start of the occupation — c. 2300 BC — until at least the early period following the deconstruction event, c. 2050 BC. While the spatial organisation and the solar alignment of the main entrances were maintained throughout the main occupation, stone axes and ‘formal’ graves indicate the continuation of the spatial concepts described above until the twentieth to nineteenth centuries BC.
These layers of meaning mirror parallel concepts of space including, although not necessarily restricted to, the formation of group identities (see Hansen & Meyer 2013: 5). They can perhaps be better understood as a ‘cosmological geography’ manifested in the symbolism of superimposed levels of conceptual ideas related to space and to certain cardinal points (Figure 8). This idea is closely related to Eliade’s (1959: 29–36) understanding of “organized — hence comicized — territory”, that is territory consecrated to provide orientation within the homogeneity of the chaotic ‘outside world’, and the equivalence of spatial consecration and cosmogony. Put differently, the Pömmelte enclosure can be interpreted as a man-made metaphor and an icon of the cosmos, reflecting the Weltanschauung (a comprehensive conception of the world) of the people who built and used it. By bringing together Eliade and Rappaport’s ideas of meaningfulness in relation to religious experience (Rappaport 1999: 391–95), it may be argued that Pömmelte was a place intended to induce oneness with the cosmos. In combining multiple layers that symbolically represent different aspects of life (first-ordermeaning), the enclosure became an icon metaphorically representing the world (second-order-meaning). As this icon was the place to reaffirm life symbolism ritually, through their actions, people perhaps experienced a sense of rootedness in, or unity with, the cosmos (highest-order-meaning). Although we can only speculate about the perceptions of ancient people, such a theory aiming to describe general principles of religious experience can provide insight.
The circular enclosure of Pömmelte is the first Central European monumental complex of primarily sacred importance that has been excavated and studied in detail. It reveals aspects of society and belief during the transition from the Final Neolithic to the Early Bronze Age, in the second half of the third millennium BC. Furthermore, it offers details of ritual behaviour and the way that people organised their landscape. A sacred interior was separated from the profane environment, and served as a venue for rites that secured the continuity of the social, spiritual and cosmic order. Ancestor worship formed another integral part of this: a mound-covered burial hut and a square-shaped ditch sanctuary (located, respectively, within and near the enclosure’s south-eastern sector; cf. Figure 2)—dating to 2880–2580 cal BC and attributed to the Corded Ware Culture (Spatzier 2017a: 235–44)—suggest that this site was deliberately chosen. With construction of the ring sanctuary, this place gained an immense expansion in meaning—comparable to Stonehenge. Through architectural transformation, both of these sites developed into sanctuaries with increasingly complex religious functions, including in relation to the cult of the dead. The cosmological and social functions, and the powerful symbolism of the Nebra sky disc and hoard (Meller 2010: 59–70), are reflected in Pömmelte’s monumental architecture.
All of these features—along with Pömmelte’s dating, function and complex ring structure—are well documented for British henge monuments (Harding 2003; Gibson 2005). The continuous use of circular enclosures in Central Europe from around 3000– 1500 BC remains to be confirmed, but strong evidence indicates usage spanning from the fifth to the first millennia BC (Spatzier 2017a: 273–96). From 2500 BC onwards, examples in Central Europe, Iberia and Bulgaria (Bertemes 2002; Escudero Carrillo et al. 2017) suggest a Europe-wide concept of sanctuary. This indicates that in extensive communication networks at the beginning of bronze metallurgy (Bertemes 2016), intellectual and religious contents circulated alongside raw materials. The henge monuments of the British Isles are generally considered to represent a uniquely British phenomenon, unrelated to Continental Europe; this position should now be reconsidered. The uniqueness of Stonehenge lies, strictly speaking, with its monumental megalithic architecture.
The Classical Bell Beaker heritage
No serious scholar can argue at this point against the male-biased East Bell Beaker migrations that expanded the European languages related to Late Proto-Indo-European-speaking Yamna (see David Reich’s comments), and thus most likely North-West Indo-European – the ancestor of Italo-Celtic, Germanic, and Balto-Slavic, apart from Pre-Celtic IE in the British Isles, Lusitano-Galician in Iberia, or Messapic in Italy (see here a full account).
With language, these migrants (several ten thousands) brought their particular Weltanschauung to all of Western, Central, and Northern Europe. Their admixture precisely in Hungary shows that they had close interactions with non-Indo-European peoples (genetically related to the Globular Amphorae culture), something that we knew from the dozens of non-Indo-European words reconstructed exclusively for North-West Indo-European, apart from the few reconstructed non-Indo-European words that NWIE shares with Palaeo-Balkan languages, which point to earlier loans from their ancestors, Yamna settlers migrating along the lower Danube.
It is not difficult to imagine that the initial East Bell Beaker group shared a newly developed common cosmological point of view that clashed with other neighbouring Yamna-related worldviews (e.g. in Balkan EBA cultures) after the cultural ties with Yamna were broken. Interesting in this respect is for example their developed (in mythology as in the new North-West Indo-European concept) *Perkwūnos, the weather god – probably remade (in language as in concept) from a Yamna minor god also behind Old Indian parjányas, the rain god – as one of the main gods from the new Pantheon, distinct from *Dyēus patēr, the almighty father sky god. In support of this, the word *meldh-n- ‘lightning’, behind the name of the mythological hammer of the weather god (cf. Old Norse Mjǫllnir or Latvian Milna), was also a newly coined North-West Indo-European term, although the myth of the hero slaying the dragon with the magical object is older.
Circular enclosures are known in Europe since the Neolithic. Also, the site selected for the Pömmelte enclosure had been used to bury Corded Ware individuals some centuries before its construction, and Corded Ware symbolism (stone axe vs. quern) is seen in the use given by Bell Beakers and later Únětice at this place. All this and other regional similarities between Bell Beakers and different local cultures (see here an example of Iberian Bell Beakers) points to syncretism of the different Bell Beaker groups with preceding cultures in the occupied regions. After all, their genealogical ancestors included also those of their maternal side, and not all encountered males disappeared, as is clearly seen in the resurge of previous paternal lineages in Central-East Europe and in Scandinavia. The admixture of Bell Beakers with previous groups (especially those of similar steppe-related ancestry from Corded Ware) needs more complex analyses to clarify potential early dialectal expansions (read what Iosif Lazaridis has to say).
The popular “big and early” expansions
These syncretic trends gave rise to distinct regional cultures, and eventually different local groups rose to power in the new cultural regions and ousted the old structures. Social norms, hierarchy, and pantheons were remade. Events like this must have been repeated again and again in Bronze and Iron Age Europe, and in many cases it was marked by a difference in the prevailing archaeological culture attested, and probably accompanied by certain population replacements that will be seen with more samples and studies of fine-scale population structure.
Some of these cultural changes, marked by evident haplogroup or admixture replacement, are defined as a ‘resurge’ of ancestry linked to previous populations, although that is obviously not equivalent to a resurge of a previous cultural group, because they usually represent just a successful local group of the same supraregional culture with a distinct admixture and/or haplogroup (see e.g. resurge of R1a-Z645 in Central-East European Bronze Age). Social, religious, or ethnic concepts may have changed in each of these episodes, along with the new prestige dialect.
This must have happened then many times during the hundreds (or thousands in some cases) of years until the first attestation of a precise ancient language and culture (read e.g. about one of the latest branches to be attested, Balto-Slavic). Ancient language contacts, like substrates or toponymy, can only rarely be detected after so many changes, so their absence (or the lack of proper studies on them) is usually not relevant – and certainly not an argument – in scholarly discussions. Their presence, on the other hand, is a proof of such contacts.
We have dozens of papers supporting Uralic dialectal substrate influence on Pre-Germanic, Proto-Balto-Slavic, and Pre- and Proto-Indo-Iranian (and even Proto-Celtic), as well as superstrate influence of Palaeo-Germanic (i.e. from Pre- to Proto-Germanic) and Proto-Balto-Slavic into Proto-Finno-Saamic, much stronger than the Indo-Iranian adstrate influence on Finno-Ugric (see the relative importance of each influence) which locates all these languages and their evolution to the north and west of the steppe (with Proto-Permic already separated, in North-East Europe, as is Proto-Ugric further east near the Urals), probably around the Baltic and Scandinavia after the expansion of Bell Beakers. These connections have been known in linguistics for decades.
Apart from some early 20th century scholars, only a minority of Indo-Europeanists support nowadays an Indo-European (i.e. centum) substrate for Balto-Slavic, to keep alive an Indo-Slavonic group based on a hypothetical 19th century Satem group; so e.g. Holzer with his Temematic, and Kortlandt supporting him, also with some supposed Indo-European substrate with heavy non-Indo-European influence for Germanic and Balto-Slavic, that now (thanks mainly to the views of the Copenhagen group) have been linked to the Corded Ware culture, as it has become clear even to them that Bell Beakers expanded North-West Indo-European.
For their part, only a minority among Uralicists, such as Kuz’mina, Parpola or Häkkinen, believe in an ‘eastern’ origin of Uralic languages, around the Southern Urals. Genomic finds – like their peers – are clearly not supporting their views. But even if we accept this hypothesis, there is little space beyond Abashevo and related East Corded Ware cultures after the recent papers on Corded Ware and Fennoscandian samples. And yet here we are:
substitutes arrows for Kron-like colors (where danger red = Indo-European) with the same end result of many other late 20th century whole-Europe Kurgan maps, linking Sredni Stog and Corded Ware with Yamna, but obviating the precise origin of Corded Ware peoples (is it Sredni Stog, or is it that immutable Middle Dnieper group? is it West Yamna, or Yamna Hungary? is it wool, or is it wheels?);
relegates Uralic speakers to a tiny corner, a ‘Volosovo’ cultural region, thus near Khvalynsk/Yamna (but not too much), that miraculously survives surrounded by all-early-splitting, all-Northern Eneolithic Indo-Europeans, thus considering Uralic languages irrelevant not only to locate the PIE Urheimat, but also to locate their own homeland; also, cultures identified in color with Uralic speakers expand until the Iron Age with enough care not to even touch in the map one of the known R1a samples published to date (because, for some people, apparently R1a must be Indo-European); and of course N1c or Siberian ancestry are irrelevant, too;
and adds findings of wheels and wool probably in support of some new ideas based on yet another correlation = causation argument (that I cannot then properly criticize without access to its reasoning beyond cute SmartArt-like symbols) similar to their model – already becoming a classic example of wrong use of statistical methods – based on the infamously named Yamnaya ancestral component™, which is obviously still used here, too.
The end result is thus similar to any other simplistic 1990s Gimbutas (or rather the recently radicalized IE Sredni Stog -> Corded Ware -> BBC version by the Danish workgroup) + 2000s R1a-map + 2010s Yamnaya ancestry™; but, hard to believe, it is published in mid-2018. A lot of hours of senseless effort, because after its publication it becomes ipso facto outdated.
For comparison of Yamna and Bell Beaker expansions, here is a recent simplistic, static (and yet more accurate) pair of maps, from the Reich Lab:
If the Copenhagen group keeps on pushing Gimbutas’ long ago outdated IE Sredni Stog -> Corded Ware theory as modified by Kristiansen, with their recently invented Corded Ware -> Bell Beaker model in genetics, at some point they are bound to clash with the Reich-Jena team, which seems to have less attachment to the classic Kurgan model and the wrong interpretations of the 2015 papers, and that would be something to behold. Because, as Cersei would say: “When you play the game of thrones, you win or you die. There is no middle ground.” And when you play the game of credibility, after so many, so wrong publications, well…
NOTE. I have been working on a similar GIS tool for quite some time, using my own maps and compiled genetic data, which I currently only use for my 2018 revision of the Indo-European demic diffusion model. Maybe within some weeks or months I will be able to publish the maps properly, after the revised papers. It’s a pitty that so much work on GIS and analysis with genetic data and cultural regions has to be duplicated, but I intend to keep some decent neutrality in my revised cultural maps, and this seems impossible at this point with some workgroups who have put all their eggs in one broken basket…
Several Iberian scholars have referred to stab-and-drag designs in both Bell-Beaker and Bronze Age ceramics (Maluquer de Motes 1956, 180, 196; Fernández-Posse 1982, 137), although these have not always been correctly appraised. In the 1980s it was finally realized that the sherds retrieved at the Boquique Cave should be dated to the Middle-Late Neolithic (4400-3300 BC), and that the same technique was also widely used in the Late Bronze Age (Fernández-Posse 1982, 147-149). Thus, nowadays it is possible to track this technique in inland Iberia at different moments throughout later prehistory (Alday and Moral 2011, 67). The earliest stab-and-drag motifs (Figure 2.2, 1) are, in fact, older than was initially thought (Fernández-Posse 1982); they actually date to the Early Neolithic (5500-4400 BC), contemporary to the Mediterranean Cardial impressed wares (Alday 2009, 135-137). There are also a few sporadic examples of stab-and-drag motifs among Bell-Beaker pottery (2600-2000 BC), such as the Ciempozuelos-style bowl from Las Carolinas (Madrid) (Figure 2.2, 2a) featuring so-called ‘symbolic’ schematic stags drawn by using this technique (Blasco and Baena 1996, 431, Lám. II; Garrido Pena 2000, 108). It is also possible to recognize this technique in a large Beaker from Molino Sanchón II (Zamora) (Abarquero et al. 2012, 206, fig. 190; Guerra-Doce et al. 2011, 812) (Figure 2.2, 2b) and there are other possible cases (e.g. Montero and Rodríguez 2008, 166, Lám. IX). Finally, the widespread use of this technique occurred in the Late Bronze Age (Figure 2.2, 3a & 3b) from c.1450 BC (e.g. Rodríguez Marcos 2007, 362-364; Abarquero 2005).
Analogies between Bell-Beaker and Bronze Age wares
Several Bell-Beaker styles can be discerned in the Iberian Meseta (e.g. Harrison 1977, 55-67; Garrido Pena 2000; 2014). In this subsection attention will be drawn primarily to the most frequent of these variants, the Ciempozuelos style, although more localised similarities can be recognised between the Beaker impressed-comb style and some early Cogotas I pottery. The Ciempozuelos ware (Delibes 1977; Harrison 1977, 19-20; Blasco 1994; Garrido Pena 2000, 116-126; Rodríguez Marcos 2007, 252-256) was widespread throughout the Meseta between 2600-2000 BC, in the same region subsequently occupied by Cogotas I communities (1800-1150 BC) (Fernández-Posse 1998; Abarquero 2005) (Figure 2.1). There is a wide array of resemblances between both pottery assemblages, a point that has been highlighted since the 1920s (e.g. Almagro Basch 1939, 143-144; Maluquer de Motes 1956, 196; Harrison 1977, 20; Jimeno 1984, 117-118).
The key ornamental traits that define the Ciempozuelos style are also reproduced among Cogotas I ware and are the following:
a) Widespread deployment among the early Cogotas I pottery of the more ubiquitous incised motifs in the Ciempozuelos style: herringbones, spikes and reticulates (Garrido Pena 2000, 119-120, fig. 48, themes 6 and 9; Rodríguez Marcos 2012, 155). During the Middle Bronze Age other less frequent themes are also similar to Bell-Beaker decorations, such as incised triangles filled with lines. Late Bronze Age wares feature the so-called ‘pseudo-Kerbschnitt’ (Rodríguez Marcos 2007, 369) which has striking precedents among Ciempozuelos ware (Harrison 1977, 20; Garrido Pena 2000, 120, fig. 48, theme 12) (Figure 2.3, 1a & 1b).
b) The extensive use of internal rim decoration, almost always deploying chevron motifs. This is ‘a Ciempozuelos leitmotiv’ (Harrison 1977, 20) in the Northern Meseta, where between 30% – 50% of all rims exhibit such a feature (Delibes 1977; Garrido Pena 2000, 163). The decoration of internal rims is even more widespread among Cogotas I vessels (Jimeno 1984; Rodríguez Marcos 2012, 158) (Figure 2.3, 1a).
c) White paste rubbed into the geometric decorations (Delibes 1977; Harrison 1977, 20; Jimeno 1984). Maluquer de Motes (1956, 186) in fact regarded excised and stab-and-drag techniques not as decorations per se, but as a way of anchoring encrusted inlays. He also reported that the bulk of rims in Cogotas I vessels exhibit white accretions (Maluquer de Motes 1956, 192) (Figure 2.3).
In addition, several authors agree on the likeness between the Bell-Beaker impressed-comb style and certain Cogotas I local pottery variants corresponding to its earliest phase (1800-1450 BC) (Garrido Pena 2000, 113-116). This is particularly striking for one micro-style from the western Meseta region, whose ceramics feature numerous impressed-comb motives (e.g. Fabián 2012; Rodríguez Marcos 2012, 158).
The relevance of emulated pottery decorations
 (…) there are grounds for proffering the view that the key creative mechanism responsible for the resemblances between apparently unrelated pottery assemblages was the emulation of standalone and very apparent decorative traits. It may constitute a good case for horizontal cultural transmission predicated upon iconic resemblances between easily imitated formal traits (Knappett 2010). Instead of spontaneous and autonomous innovations, it is far more compelling to regard these decorative features as interlinked and punctuated ‘way stations along the trails of living beings, moving through a world’ (Ingold and Hallam 2007, 8). No creative act can be regarded as really isolated. Instead it ought to be understood as focusing on the nodes in particular fields of associations (Lohnmann 2010, 216).
 Pottery ornamentation in the Cogotas I tradition combined and reinterpreted both local atavistic (e.g. Abarquero 2005, 24-26; Rodríguez Marcos 2007, 357-367) and widespread pan-European ornaments (e.g. Blasco 2001, 225, 2003, 67-68; Abarquero 2012, 98-101). From a semiotic perspective such things transcended large spatio-temporal distances; they were closely associated by iconical shared links in a relational or cognitive space, whereby these entities were co-presented and indirectly recalled and perceived despite being distant (Knappett 2010, 85-86). The locally-rooted biases of these creative quotations can be glimpsed from rare sequences of ceramic productions spanning several generations of potters. For instance, at Majaladares (Borja, Zaragoza) strong analogies arise between Ciempozuelos wares featuring unique decorations in this site and Cogotas I wares from the superimposed layers, exhibiting remarkably similar themes (Harrison 2007, 65-82). Likewise, it is noteworthy that the earliest triangular excisions in Cogotas I wares occurred in the eastern Meseta, where imported Duffaits vessels featuring comparable motifs were circulating from several centuries before.(…)
 There is scope for advocating that these pottery decorations cannot be envisaged as a form of irrelevant or mundane aesthetic garnish for the sake of art. Bronze Age potters drew upon a highly meaningful array of esoteric sources and, in so doing, the vessels might have echoed designs betokening genealogical, mythical or parallel worlds, in a kind of dialectical negotiation between self and other (Taussig 1993). The very involvement of ancestors and spiritual forces in making and embellishing a pot is supported by ethnographic evidence (e.g. Crown 2007, 679; Lohnmann 2010, 222) and this also seems plausible in the case of Cogotas I ceramics. These real or imagined beings might be regarded as inspiring sources of creations, whose role is often to legitimize and guarantee the accuracy of the involved knowledge (Lohnmann 2010, 222). In the same vein, the smearing of colored inlays on certain pots ought to be properly understood beyond an aesthetic action of embellishment, as our own rationale prompts us to assume. (…)
 Furthermore, this pottery tradition needs to be understood as an effective means of socialization and a key resource in the forging of identities. Decorating certain intricate Cogotas I vessels (Figure 2.2, 3b; Figure 2.4, 3) very likely involved an ostentatious difficulty (Robb and Michelaki 2012, 168; Abarquero 2005, 438) and the proficiency displayed in such tasks may have accrued even moral connotations (Hendon 2010, 146-147). Learning to perform some of the pottery decoration discussed here certainly required complex training processes involving both expert potters and mentored apprentices (Crown 2007; Hosfield 2009, 46). Thus, the stab-and-drag technique demanded time-consuming learning as well as careful and thorough execution (Alday 2009, 11-19). Likewise the selection and processing of particular raw materials – mainly bones – to attain the white inlays involved direct observation and hands-on training (Odriozola et al. 2012, 150). (…)
 Finally, the role of the Cogotas I pottery decoration was also deeply rooted in the sphere of social interactions through particular communal practices of exhibition and consumption. The celebration of commensality rituals is very often predicated as a key social practice among these communities (e.g. Harrison 1995, 74; Abarquero 2005, 56; Blanco-González 2014, 453). Potters embodied and replicated non-discursive shared tenets on a routine basis, but by means of these social gatherings and the deployment of such festive services ‘their visual materialisation made them part of the habitus of everybody’ (Chapman and Gaydarska 2007, 182). Bronze Age groups in the Meseta have recently been characterized as scarcely integrated, short-lasting and unstable social units, lacking long-term cultural rules and institutions, restricted to one generation lifespan at the most (Blanco-González 2015). (…)
If we take into account that the earliest Iberian Bell Beakers were I2a, R1b-V88, and G2a, just like previous Chalcolithic and Neolithic Iberians, it cannot get clearer how and when the first Indo-European waves reached Iberia, and thus that the Harrison and Heyd (2007) model of East Bell Beaker expansion was right. Not a single reputable geneticist contests the origin of R1b-L23 subclades in Iberia anymore (see e.g. Heyd, or Lazaridis).
While the Spanish archaeological school will be slow to adapt to genetic finds – since there are many scholars who have supported for years other ways of expansion of the different Bell Beaker motifs, and follow mostly the “pots not people” descriptive Archaeology – , many works like these can be just as well reinterpreted in light of what we already know happened in terms of population movements during this period, and this alone gives a whole new interesting perspective to archaeological finds.
On the previous, non-Indo-European stage of the Iberian Paeninsula, there is also a new paper (behind paywall), showing reasons for inter-regional differences, and thus supporting homogeneity before the arrival of Bell Beakers:
The Chalcolithic period is traditionally defined by the emergence of copper elements and associated to the beginning of defensive-style architecture (Esquivel and Navas 2007). This last characteristic only seems to appear clearly in the southeast of the Iberian Peninsula, with the denominated Millares Culture (e.g. García Sanjuán 2013; Valera et al. 2014). In the rest of the Iberian Peninsula, the Neolithic-Chalcolithic transition is scarcely defined. In fact, it is possible that this transition does not even strictly exist and rather results from the evolution of villages present in the most advanced phases of the Neolithic (e.g. Blasco et al. 2007). This continuity is also perceptible in most of the sepulchral caves over time, where radiocarbon dates show a continued use from the 4th to the 3rd millennium cal B.C. (Fernández-Crespo 2016; Utrilla et al. 2015; Villalba-Mouco et al. 2017). Moreover, it is possible to find some copper materials normally associated with burial contexts as prestigious grave goods (Blasco and Ríos 2010), but not as evidence of a massive replacement of commonly used tools such as flint blades, bone industry, polished stones or pottery without singular characteristics from a unique period (Pérez-Romero et al. 2017). (…)
The human isotope values from both sites portray a quite homogeneous overall diet among humans. This homogeneous pattern of diet based on C3 terrestrial resources seems to be general along the entire Iberian Peninsula during the Late Neolithic and Chalcolithic (e.g. Alt et al. 2016; Díaz-Zorita 2014; Fernández-Crespo et al. 2016; Fontanals-Coll et al. 2015; García-Borja et al. 2013; López-Costas et al. 2015; McClure et al. 2011; Sarasketa-Gartzia et al. 2017; Villalba- Mouco et al. 2017; Salazar-García 2011; Salazar-García et al. 2013b; Salazar-García 2014; Waterman et al. 2016). The reason of this homogeneity could be the consolidated economy based on agriculture and livestock, together with a higher mobility among the different communities and the increase of trade networks, not only in prestigious objects (Schuhmacher and Banerjee 2012) but also in food products. Isotopic analyses in fauna remains could give us more clues about animal trade, as happens in other chronologies (Salazar- García et al. 2017).
In any case, and even if the dietary interpretation does not vary, it is noteworthy to mention that there are significant differences between δ13C human values from Cova de la Guineu and δ13C human values from Cueva de Abauntz (Mann-Whitney test, p = 1.05× 10−12) (Fig. 6). This observed δ13C differences among humans is also present among herbivores (Mann-Whitney test, p = 0.0004), which define the baseline of each ecosystem. This suggests that the observed human difference between sites should not be attributed to diet, but most possibly to the existence of enough environmental differences to be recorded in the collagen δ13C values along the food web. Plants are very sensitive to different environmental factors (altitude, temperature, luminosity or water availability) and their physiological adaptation to its factors can generate a variation in their isotopic values as happens with C3 and C4 adaptations (O’Leary 1981; Ambrose 1991). This spectrum of values has been used to assess several aspects about past environmental conditions when studying the δ13C and δ15N isotopic values of a species with a fixed diet over time (e.g. Stevens et al. 2008; González-Guarda et al. 2017). Moreover, this gradual δ13C and δ15N variation among different environments is very helpful to discriminate altitudinal movements in herbivores with a high precision method based on serial dentine analysis (Tornero et al. 2016b). In our case, results reflect the influence of environment from at least two areas in Iberia (the Western Prepyrenees and the Northeastern coast of Iberia). These differences demand caution when interpreting human diets from different sites that are not contemporary and/or not in a same area, as it is possible that the environmental influence is responsible for changes otherwise attributed to different subsistence patterns and social structures (Fernández-Crespo and Schulting 2017), as has been demonstrated in neighbouring territories (Herrscher and Bras-Goude 2010; Goude and Fontugne 2016).
The Middle Neolithic is known to mark the westward expansion of Comb Ware and related cultures in North-Eastern Europe.
Mathieson et al. (2017 and 2018) had this to say about the Middle Neolithic in the Baltic:
At Zvejnieki in Latvia, using 17 newly reported individuals and additional data for 5 previously reported34 individuals, we observe a transition in hunter-gatherer-related ancestry that is opposite to that seen in Ukraine. We find that Mesolithic and Early Neolithic individuals (labelled ‘Latvia_HG’) associated with the Kunda and Narva cultures have ancestry that is intermediate between WHG (approximately 70%) and EHG (approximately 30%), consistent with previous reports34–36(Supplementary Table 3). We also detect a shift in ancestry between Early Neolithic individuals and those associated with the Middle Neolithic Comb Ware complex (labelled ‘Latvia_MN’), who have more EHG-related ancestry; we estimate that the ancestry of Latvia_MN individuals comprises 65% EHG-related ancestry, but two of the four individuals appear to be 100% EHG in principal component space (Fig. 1b).
Other samples and errors on Y-SNP calls
The truth is, this is another sample (Latvia_MN_dup.I4627.SG) from the same individual ZVEJ26.
There is another sample used for the analysis of ZVEJ26, with the same data as in Mathieson et al. (2018), i.e. better coverage, and Y-DNA R1b1a1a(xR1b1a1a2).
Most samples in the tables from Wang et al. (2018) seem to be classified correctly, as in previous papers, but for:
Blätterhöhle Cave sample from Lipson et al. (2017), wrongly classified (again) as R1b1a1a2a1a2a1b2 (I am surprised no R1b-autochtonous-continuity-fan rushed to proclaim something based on this);
Mal’ta 1 sample from Raghavan et al. (2013) as R1b1a1a2;
Iron Gates HG, Schela Cladovey from Gonzalez Fortes (2017) as R1b1a1a2;
Oase1 from Fu (2015) as N1c1a;
samples from Skoglund et al. (2017) from Africa also wrongly classified as R1b1a1a2 and subclades.
It seems therefore that the poor coverage / SNPs hit on autosomes is the key common factor here for these Y-SNP calls, and so it is in the Zvejnieki MN1 duplicated sample. Anyway, if all Y-SNP calls come from the same software applied to all data, and this is going to be used in future papers, this seems to be a great improvement compared to Narasimhan et al. (2018)…
EDIT (25 JUN 2018): I have been reviewing some more papers apart from Mathieson et al. (2018) and Olalde et al. (2018) to compare the reported haplogroups, and there seems to be many potential errors (or updated data, difficult to say sometimes, especially when the newly reported haplogroup is just one or two subclades below the reported one in ‘old’ papers), not only those listed above.
The sample accession number in the European Nucleotide Archive (ENA) is SAMEA45565168 (Latvia_MN1/ZVEJ26) (see here), in case anyone used to this kind of analysis wishes to repeat the Y-SNP calls on both samples.
EDIT (25 JUN 2018): Added that it is another sample with lesser coverage from the same ZVEJ26 individual.
Nevertheless, because I keep reading the same comments no matter what gets published, even in mid-2018 – the latest ones in our Facebook page (“was haplogroup X Indo-European?”), and in this very blog (“I see it very difficult to link Bell Beaker with Balto-Slavic, when now Balto-Slavic people are strikingly R1a-dominated”); and because I see even more misunderstandings and personal attacks, I have decided to publish them.
This way I will be able to explain my “R1b-L23/Proto-Indo-Europeans” theory with simplistic maps (however badly I hate such maps when I find them on Google searches), and I will also have a page to redirect those who don’t want to dismiss the “R1a – Indo-European association”, instead of answering comments about this question each time they pop up…
Here you have the links to the posts – and also on the menu above (there is a lot of rambling, because they are from a period of less clear data on Yamna and Corded Ware; today I would have never written such long discussions, they are mostly unnecessary):
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.
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).
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.
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.
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 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.
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.
The most interesting of the local people is the occupant of grave 12, which is the earliest grave in the kurgan and the main statistical range of its radiocarbon date clearly predates the arrival of the western Yamnaya groups c. 3000 BC. This is also confirmed by the burial rite, which is not typical for the Yamnaya (Dani 2011: 29–33; Heyd in press), although some heterogeneity may apply in Yamnaya communities too. The migrant group, graves nos. 4, 7, 9 and 11, all occupy late stratigraphic positions in the mound, and have radiocarbon dates in the second quarter of the third millennium BC. It is also noteworthy that they are all adult or mature men. The contextual data, their physical distribution over the space of the whole kurgan, and the variety of burial practices, indicate several generations of burials. The cultural attributes of this group are summarised in Figure 5. Overall, their closest match lies in the Livezile group from the eastern and southern Apuseni Mountains, which is also the likely place of origin of the buried persons.
The key question is, what cultural process could be responsible for attracting these men from their homeland to the Great Hungarian Plain, over several generations? Their sex and age uniformity indicate they are a social sub-set within a larger group, implying that only a portion of their society was on the move. Exogamy can probably be excluded, since one would expect more women than men to move in prehistoric times; not to mention the distance of more than 200km between the places of potential origin and burial.
One hypothesis would see these men involved in the exchange of goods, with long-term relations between the mountain and steppe communities. Normally living in, or next to, the Apuseni, these men would journey for weeks into the plain, returning to the same places and people over many decades. Ethnographic examples of such travels to exchange objects and ideas, and perhaps people, are numerous (e.g. Helms 1988). However, the child’s (grave 7a) local isotopic signature would remain unexplained, and one has to wonder for how many generations an exchange continues for four men to die near the Őrhalom.
A second hypothesis is essentially an economic model of transhumance, with livestock passing the winter and spring in the milder regions of the Great Hungarian Plain, and returning to higher pastures in the warmer months (Arnold & Greenfield 2006). Such systems can endure for centuries, provided the social relations underpinning them are stable. This has the advantage of accounting for relatively long periods of time spent away from home, as herdsmen guarded their animals, and perhaps some women and their children came too, which would account for the child’s presence, and the pottery relations of the Livezile group. Furthermore, regular visits to a region would increase the likelihood of Livezile transhumant herders becoming integrated locally. The second quarter of the third millennium BC was a period when Yamnaya ideology, and thus its internal coherence, might have already diminished. This would likely have resulted in a weakened grip by Yamnaya people on pastures and territory, consequently allowing Livezile herders, and potentially others, to step in and take over locally, perhaps first on a seasonal basis and then permanently.
On West Yamna settlers in Hungary
By disclosing very interesting information on (yet unpublished) Yamna samples from Hungary, the latest preprint from the Reich Lab has rendered irrelevant – in a rather surprising turn of events – (what I expected would be) future discussions on West Yamna settlers potentially sharing a similar ancestry with Baltic Late Neolithic / Corded Ware settlers (see here for more details).
Interesting excerpts regarding the tight cluster formed by all Yamna samples:
Individuals from the North Caucasian steppe associated with the Yamnaya cultural formation (5300-4400 BP, 3300-2400 calBCE) appear genetically almost identical to previously reported Yamnaya individuals from Kalmykia20 immediately to the north, the middle Volga region19, 27,Ukraine and Hungary, and to other Bronze Age individuals from the Eurasian steppes who share the characteristic ‘steppe ancestry’ profile as a mixture of EHG and CHG/Iranian ancestry23, 28. These individuals form a tight cluster in PCA space (Figure 2) and can be shown formally to be a mixture by significantly negative admixture f3-statistics of the form f3(EHG, CHG; target) (Supplementary Fig. 3).
Using qpAdm with Globular Amphora as a proximate surrogate population (assuming that a related group was the source of the Anatolian farmer-related ancestry), we estimated the contribution of Anatolian farmer-related ancestry into Yamnaya and other steppe groups. We find that Yamnaya individuals from the Volga region (Yamnaya Samara) have 13.2±2.7% and Yamnaya individuals in Hungary 17.1±4.1% Anatolian farmer-related ancestry (Fig.4; Supplementary Table 18)– statistically indistinguishable proportions.
Before this paper, we had the solidest anthropological models backed by Y-DNA against conflicting data from certain statistical tools applied to a few samples (which some used to contradict what was mainstream in Academia).
Today, we have everything – including statistical tools – showing a genetically homogeneous, Late PIE-speaking late Khvalynsk/Yamna community expanding into its known branches, confirming what was described using traditional anthropological disciplines:
Late Khvalynsk expanding into Afanasevo ca. 3300-3000 BC with an archaic Late PIE dialect, which was attested much later as Tocharian;
and now also Yamna settlers: those in Hungary admixing (probably ca. 2800-2500 BC) with the local population to form North-West Indo-European-speaking East Bell Beakers; those from the Balkans forming other IE-speaking Balkan cultures, including the peoples that admixed in Greece, as seen in Mycenaeans.