Vikings, Vikings, Vikings! “eastern” ancestry in the whole Baltic Iron Age

vikings-middle-age

Open access Population genomics of the Viking world, by Margaryan et al. bioRxiv (2019), with a huge new sampling from the Viking Age.

Interesting excerpts (emphasis mine, modified for clarity):

To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago.

Maps illustrating the following texts have been made based on data from this and other papers:

  • Maps showing ancestry include only data from this preprint (which also includes some samples from Sigtuna).
  • Maps showing haplogroup density include Vikings from other publications, such as those from Sigtuna in Krzewinska et al. (2018), and from Iceland in Ebenesersdóttir et al. (2018).
  • Maps showing haplogroups of ancient DNA samples based on their age include data from all published papers, but with slightly modified locations to avoid overcrowding (randomized distance approx. ± 0.1 long. and lat.).

middle-ages-europe-y-dna
Y-DNA haplogroups in Europe during the Viking expansions (full map). See other maps from the Middle Ages.

We find that the transition from the BA to the IA is accompanied by a reduction in Neolithic farmer ancestry, with a corresponding increase in both Steppe-like ancestry and hunter-gatherer ancestry. While most groups show a slight recovery of farmer ancestry during the VA, there is considerable variation in ancestry across Scandinavia. In particular, we observe a wide range of ancestry compositions among individuals from Sweden, with some groups in southern Sweden showing some of the highest farmer ancestry proportions (40% or more in individuals from Malmö, Kärda or Öland).

Ancestry proportions in Norway and Denmark on the other hand appear more uniform. Finally we detect an influx of low levels of “eastern” ancestry starting in the early VA, mostly constrained among groups from eastern and central Sweden as well as some Norwegian groups. Testing of putative source groups for this “eastern” ancestry revealed differing patterns among the Viking Age target groups, with contributions of either East Asian- or Caucasus-related ancestry.

saami-ancestry-vikings
Ancestry proportions of four-way models including additional putative source groups for target groups for which three-way fit was rejected (p ≤ 0.01);

Overall, our findings suggest that the genetic makeup of VA Scandinavia derives from mixtures of three earlier sources: Mesolithic hunter-gatherers, Neolithic farmers, and Bronze Age pastoralists. Intriguingly, our results also indicate ongoing gene flow from the south and east into Iron Age Scandinavia. Thus, these observations are consistent with archaeological claims of wide-ranging demographic turmoil in the aftermath of the Roman Empire with consequences for the Scandinavian populations during the late Iron Age.

Genetic structure within Viking-Age Scandinavia

We find that VA Scandinavians on average cluster into three groups according to their geographic origin, shifted towards their respective present-day counterparts in Denmark, Sweden and Norway. Closer inspection of the distributions for the different groups reveals additional complexity in their genetic structure.

vikings-danish-ancestry
Natural neighbor interpolation of “Danish ancestry” among Vikings.

We find that the ‘Norwegian’ cluster includes Norwegian IA individuals, who are distinct from both Swedish and Danish IA individuals which cluster together with the majority of central and eastern Swedish VA individuals. Many individuals from southwestern Sweden (e.g. Skara) cluster with Danish present-day individuals from the eastern islands (Funen, Zealand), skewing towards the ‘Swedish’ cluster with respect to early and more western Danish VA individuals (Jutland).

Some individuals have strong affinity with Eastern Europeans, particularly those from the island of Gotland in eastern Sweden. The latter likely reflects individuals with Baltic ancestry, as clustering with Baltic BA individuals is evident in the IBS-UMAP analysis and through f4-statistics.

vikings-norwegian-ancestry
Natural neighbor interpolation of “Norwegian ancestry” among Vikings.

For more on this influx of “eastern” ancestry see my previous posts (including Viking samples from Sigtuna) on Genetic and linguistic continuity in the East Baltic, and on the Pre-Proto-Germanic homeland based on hydrotoponymy.

Baltic ancestry in Gotland

Genetic clustering using IBS-UMAP suggested genetic affinities of some Viking Age individuals with Bronze Age individuals from the Baltic. To further test these, we quantified excess allele sharing of Viking Age individuals with Baltic BA compared to early Viking Age individuals from Salme using f4 statistics. We find that many individuals from the island of Gotland share a significant excess of alleles with Baltic BA, consistent with other evidence of this site being a trading post with contacts across the Baltic Sea.

vikings-finnish-ancestry
Natural neighbor interpolation of “Finnish ancestry” among Vikings.

The earliest N1a-VL29 sample available comes from Iron Age Gotland (VK579) ca. AD 200-400 (see Iron Age Y-DNA maps), which also proves its presence in the western Baltic before the Viking expansion. The distribution of N1a-VL29 and R1a-Z280 (compared to R1a in general) among Vikings also supports a likely expansion of both lineages in succeeding waves from the east with Akozino warrior-traders, at the same time as they expanded into the Gulf of Finland.

vikings-y-dna-haplogroup-r1a-z280-over-r1a
Density of haplogroup R1a-Z280 (samples in pink) overlaid over other R1a samples (in green, with R1a-Z284 in cyan) among Vikings.

Vikings in Estonia

(…) only one Viking raiding or diplomatic expedition has left direct archaeological traces, at Salme in Estonia, where 41 Swedish Vikings who died violently were buried in two boats accompanied by high-status weaponry. Importantly, the Salme boat-burial predates the first textually documented raid (in Lindisfarne in 793) by nearly half a century. Comparing the genomes of 34 individuals from the Salme burial using kinship analyses, we find that these elite warriors included four brothers buried side by side and a 3rd degree relative of one of the four brothers. In addition, members of the Salme group had very similar ancestry profiles, in comparison to the profiles of other Viking burials. This suggests that this raid was conducted by genetically homogeneous people of high status, including close kin. Isotope analyses indicate that the crew descended from the Mälaren area in Eastern Sweden thus confirming that the Baltic-Mid-Swedish interaction took place early in the VA.

vikings-swedish-ancestry
Natural neighbor interpolation of “Swedish ancestry” among Vikings.

Viking samples from Estonia show thus ancient Swedes from the Mälaren area, which proves once again that hg. N1a-VL29 (especially subclade N1a-L550) and tiny proportions of so-called “Siberian ancestry” expanded during the Early Iron Age into the whole Baltic Sea area, not only into Estonia, and evidently not spreading with Balto-Finnic languages (since the language influence is in the opposite direction, east-west, Germanic > Finno-Samic, during the Bronze Age).

N1a-VL29 lineages spread again later eastwards with Varangians, from Sweden into north-eastern Europe, most likely including the ancestors of the Rurikid dynasty. Unsurprisingly, the arrival of Vikings with Swedish ancestry into the East Baltic and their dispersal through the forest zone didn’t cause a language shift of Balto-Finnic, Mordvinic, or East Slavic speakers to Old Norse, either…

NOTE. For N1a-Y4339 – N1a-L550 subclade of Swedish origin – as main haplogroup of modern descendants of Rurikid princes, see Volkov & Seslavin (2019) – full text in comments below. Data from ancient samples show varied paternal lineages even among early rulers traditionally linked to Rurik’s line, which explains some of the discrepancies found among modern descendants:

  • A sample from Chernihiv (VK542) potentially belonging to Gleb Svyatoslavich, the 11th century prince of Tmutarakan/Novgorod, belongs to hg. I2a-Y3120 (a subclade of early Slavic I2a-CTS10228) and has 71% “Modern Polish” ancestry (see below).
  • Izyaslav Ingvarevych, the 13th century prince of Dorogobuzh, Principality of Volhynia/Galicia, is probably behind a sample from Lutsk (VK541), and belongs to hg. R1a-L1029 (a subclade of R1a-M458), showing ca. 95% of “Modern Polish” ancestry.
  • Yaroslav Osmomysl, the 12th century Prince of Halych (now in Western Ukraine), was probably of hg. E1b-V13, yet another clearly early Slavic haplogroup.

vikings-y-dna-haplogroup-n1a
Density of haplogroup N1a-VL29, N1a-L550 (samples in pink, most not visible) among Vikings. Samples of hg. R1b in blue, hg. R1a in green, hg. I in orange.

Finnish ancestry

Firstly, modern Finnish individuals are not like ancient Finnish individuals, modern individuals have ancestry of a population not in the reference; most likely Steppe/Russian ancestry, as Chinese are in the reference and do not share this direction. Ancient Swedes and Norwegians are more extreme than modern individuals in PC2 and 4. Ancient UK individuals were more extreme than Modern UK individuals in PC3 and 4. Ancient Danish individuals look rather similar to modern individuals from all over Scandinavia. By using a supervised ancient panel, we have removed recent drift from the signal, which would have affected modern Scandinavians and Finnish populations especially. This is in general a desirable feature but it is important to check that it has not affected inference.

ancient-modern-finns-steppe
PCA of the ancient and modern samples using the ancient palette, showing different PCs. Modern individuals are grey and the K=7 ancient panel surrogate populations are shown in strong colors, whilst the remaining M-K=7 ancient populations are shown in faded colors.

The story for Modern-vs-ancient Finnish ancestry is consistent, with ancient Finns looking much less extreme than the moderns. Conversely, ancient Norwegians look like less-drifted modern Norwegians; the Danish admixture seen through the use of ancient DNA is hard to detect because of the extreme drift within Norway that has occurred since the admixture event. PC4 vs PC5 is the most important plot for the ancient DNA story: Sweden and the UK (along with Poland, Italy and to an extent also Norway) are visibly extremes of a distribution the same “genes-mirror-geography” that was seen in the Ancient-palette analysis. PC1 vs PC2 tells the same story – and stronger, since this is a high variance-explained PC – for the UK, Poland and Italy.

Uniform manifold approximation and projection (UMAP) analysis of the VA and other ancient samples.

Evidence for Pictish Genomes

The four ancient genomes of Orkney individuals with little Scandinavian ancestry may be the first ones of Pictish people published to date. Yet a similar (>80% “UK ancestry) individual was found in Ireland (VK545) and five in Scandinavia, implying that Pictish populations were integrated into Scandinavian culture by the Viking Age.

Our interpretation for the Orkney samples can be summarised as follows. Firstly, they represent “native British” ancestry, rather than an unusual type of Scandinavian ancestry. Secondly, that this “British” ancestry was found in Britain before the Anglo-Saxon migrations. Finally, that in Orkney, these individuals would have descended from Pictish populations.

vikings-british-ancestry
Natural neighbor interpolation of “British ancestry” among Vikings.

(…) ‘UK’ represents a group from which modern British and Irish people all receive an ancestry component. This information together implies that within the sampling frame of our data, they are proxying the ‘Briton’ component in UK ancestry; that is, a pre-Roman genetic component present across the UK. Given they were found in Orkney, this makes it very likely that they were descended from a Pictish population.

Modern genetic variation within the UK sees variation between ‘native Briton’ populations Wales, Scotland, Cornwall and Ireland as large compared to that within the more ‘Anglo-Saxon’ English. This is despite subsequent gene flow into those populations from English-like populations. We have not attempted to disentangle modern genetic drift from historically distinct populations. Roman-era period people in England, Wales, Ireland and Scotland may not have been genetically close to these Orkney individuals, but our results show that they have a shared genetic component as they represent the same direction of variation.

Density of haplogroup R1b-L21 (samples in red), overlaid over all samples of hg. R1b among Vikings (R1b-U106 in green, other R1b-L151 in deep red). To these samples one may add the one from Janakkala in south-western Finland (AD ca. 1300), of hg. R1b-L21, possibly related to these population movements.

For more on Gaelic ancestry and lineages likely representing slaves among early Icelanders, see Ebenesersdóttir et al. (2018).

Y-DNA

As in the case of mitochondrial DNA, the overall distribution profile of the Y chromosomal haplogroups in the Viking Age samples was similar to that of the modern North European populations. The most frequently encountered male lineages were the haplogroups I1, R1b and R1a.

Haplogroup I (I1, I2)

The distribution of I1 in southern Scandinavia, including a sample from Sealand (VK532) ca. AD 100 (see Iron Age Y-DNA maps) proves that it had become integrated into the West Germanic population already before their expansions, something that we already suspected thanks to the sampling of Germanic tribes.

vikings-y-dna-haplogroup-i
Density of haplogroup I (samples in orange) among Vikings. Samples of hg. R1b in blue, hg. R1a in green, N1a in pink.
vikings-y-dna-haplogroup-i1-over-i
Density of haplogroup I1 (samples in red) overlaid over all samples of hg. I among Vikings.

Haplogroup R1b (M269, U106, P312)

Especially interesting is the finding of R1b-L151 widely distributed in the historical Nordic Bronze Age region, which is in line with the estimated TMRCA for R1b-P312 subclades found in Scandinavia, despite the known bottleneck among Germanic peoples under U106. Particularly telling in this regard is the finding of rare haplogroups R1b-DF19, R1b-L238, or R1b-S1194. All of that points to the impact of Bell Beaker-derived peoples during the Dagger period, when Pre-Proto-Germanic expanded into Scandinavia.

Also interesting is the finding of hg. R1b-P297 in Troms, Norway (VK531) ca. 2400 BC. R1b-P297 subclades might have expanded to the north through Finland with post-Swiderian Mesolithic groups (read more about Scandinavian hunter-gatherers), and the ancestry of this sample points to that origin.

However, it is also known that ancestry might change within a few generations of admixture, and that the transformation brought about by Bell Beakers with the Dagger Period probably reached Troms, so this could also be a R1b-M269 subclade. In fact, the few available data from this sample show that it comes from the natural harbour Skarsvågen at the NW end of the island Senja, and that its archaeologist thought it was from the Viking period or slightly earlier, based on the grave form. From Prescott (2017):

In 1995, Prescott and Walderhaug tentatively argued that a dramatic transformation took place in Norway around the Late Neolithic (2350 BCE), and that the swift nature of this transition was tied to the initial Indo-Europeanization of southern and coastal Norway, at least to Trøndelag and perhaps as far north as Troms. (…)

The Bell Beaker/early Late Neolithic, however, represents a source and beginning of these institution and practices, exhibits continuity to the following metal age periods and integrated most of Northern Europe’s Nordic region into a set of interaction fields. This happened around 2400 BCE, at the MNB to LN transition.

NOTE. This particular sample is not included in the maps of Viking haplogroups.

vikings-y-dna-haplogroup-r1b
Density of haplogroup R1b (samples in blue) among Vikings. Samples of hg. I in orange, hg. R1a in green, N1a in pink.
vikings-y-dna-haplogroup-r1b-U106-over-r1b
Density of haplogroup R1b-U106 (samples in green) overlaid over all samples of hg. R1b (other R1b-L23 samples in red) among Vikings.
vikings-y-dna-haplogroup-r1b-P312-over-r1b
Density of R1b-L151 (xR1b-U106) (samples in deep red) overlaid over all samples of hg. R1b (R1b-U106 in green, other R1b-M269 in blue) among Vikings.

Haplogroup R1a (M417, Z284)

The distribution of hg. R1a-M417, in combination with data on West Germanic peoples, shows that it was mostly limited to Scandinavia, similar to the distribution of I1. In fact, taking into account the distribution of R1a-Z284 in particular, it seems even more isolated, which is compatible with the limited impact of Corded Ware in Denmark or the Northern European Plain, and the likely origin of R1a-Z284 in the expansion with Battle Axe from the Gulf of Finland. The distribution of R1a-Z280 (see map above) is particularly telling, with a distribution around the Baltic Sea mostly coincident with that of N1a.

vikings-y-dna-haplogroup-r1a
Density of haplogroup R1a (samples in green) among Vikings. Samples of hg. R1b in blue, of hg. I in orange, N1a in pink.
vikings-y-dna-haplogroup-r1a-z284-over-r1a
Density of haplogroup R1a-Z284 (samples in cyan) overlaid over all samples of hg. R1a (in green, with R1a-Z280 in pink) among Vikings.

Other haplogroups

Among the ancient samples, two individuals were derived haplogroups were identified as E1b1b1-M35.1, which are frequently encountered in modern southern Europe, Middle East and North Africa. Interestingly, the individuals carrying these haplogroups had much less Scandinavian ancestry compared to the most samples inferred from haplotype based analysis. A similar pattern was also observed for less frequent haplogroups in our ancient dataset, such as G (n=3), J (n=3) and T (n=2), indicating a possible non-Scandinavian male genetic component in the Viking Age Northern Europe. Interestingly, individuals carrying these haplogroups were from the later Viking Age (10th century and younger), which might indicate some male gene influx into the Viking population during the Viking period.

vikings-italian-ancestry
Natural neighbor interpolation of “Italian ancestry” among Vikings.

As the paper says, the small sample size of rare haplogroups cannot distinguish if these differences are statistically relevant. Nevertheless, both E1b samples have substantial Modern Polish-like ancestry: one sample from Gotland (VK474), of hg. E1b-L791, has ca. 99% “Polish” ancestry, while the other one from Denmark (VK362), of hg. E1b-V13, has ca. 35% “Polish”, ca. 35% “Italian”, as well as some “Danish” (14%) and minor “British” and “Finnish” ancestry.

Given the E1b-V13 samples of likely Central-East European origin among Lombards, Visigoths, and especially among Early Slavs, and the distribution of “Polish” ancestry among Viking samples, VK362 is probably a close description of the typical ancestry of early Slavs. The peak of Modern Polish-like ancestry around the Upper Pripyat during the (late) Viking Age suggests that Poles (like East Slavs) have probably mixed since the 10th century with more eastern peoples close to north-eastern Europeans, derived from ancient Finno-Ugrians:

vikings-polish-ancestry
Natural neighbor interpolation of “Polish ancestry” among Vikings.

Similarly, the finding of R1a-M458 among Vikings in Funen, Denmark (VK139), in Lutsk, Poland (VK541), and in Kurevanikha, Russia (VK160), apart from the early Slav from Usedom, may attest to the origin of the spread of this haplogroup in the western Baltic after the Bell Beaker expansion, once integrated in both Germanic and probably Balto-Slavic populations, as well as intermediate Bronze Age peoples that were eventually absorbed by their expansions. This contradicts, again, my simplistic initial assessment of R1a-M458 expansion as linked exclusively (or even mainly) to Balto-Slavs.

antiquity-europe-y-dna
Y-DNA haplogroups in Europe during Antiquity (full map). See other maps of cultures and ancient DNA from Antiquity.

Related

Genetic continuity among Uralic-speaking cultures in north-eastern Europe

east-europe-bronze-age

The recent study of Estonian Late Bronze Age/Iron Age samples has shown, as expected, large genetic continuity of Corded Ware populations in the East Baltic area, where West Uralic is known to have been spoken since at least the Early Bronze Age.

The most interesting news was that, unexpectedly for many, the impact of “Siberian ancestry” (whatever that actually means) was small, slow, and gradual, with slight increases found up to the Middle Ages, compatible with multiple contact events in north-eastern Europe. Haplogroup N became prevalent among Finnic populations only through late bottlenecks, as research of modern populations have long suggested, and as ancient DNA research hinted since at least 2015.

I risked to correlate the arrival of chiefs from the south-west with the infiltration of N1c-VL29 subclades during the transition to the Iron Age, coupled with that minimal “Siberian” ancestry (see e.g. here and here). Now we know that the penetration of this non-CW ancestry started, as predicted, in the Iron Age; that it was highly variable in the few samples where it appeared, with ca. 1-4%, while most Iron Age individuals show 0%; and that it was not especially linked to individuals of N1c-Vl29 lineages.

It is also basically confirmed, based on the (ancient and Modern Swedish) N1c-L550 subclades found among Iron Age Estonians, that N1c-VL29 lineages and the so-called “Siberian” ancestry will be found simultaneously around the Baltic coastal areas, and that different lineages must have suffered later founder effects among Finns, which suggests that these alliances through exogamy brought exactly as much language change in Sweden, Lithuania, or Poland, as they did in the East Baltic region…

On the other hand, the paper has also shown a potential movement of Corded Ware-derived peoples, if the change from LBA to IA samples is meaningful; in fact, even more Corded Ware-like than Baltic and Estonian BA populations. The exact origin of that movement is difficult to pinpoint, and it may not be related to the arrival of Akozino warrior-traders from the south-east, since theirs seems to be a minor impact proper of elites in a chiefdom system around the Baltic.

fortified-settlements-lba-ia
Distribution of fortified settlements (filled circles) and other hilltop sites (empty circles) of the Late Bronze Age and Pre-Roman Iron Ages in the East Baltic region. Tentative area of most intensive contacts between Baltic and Balto-Finnic communities marked with a dashed line. Image modified from (Lang 2016).

Also suggesting a potential movement is the ‘southern’ shift observed in the West and East Baltic areas, likely showing the arrival of Proto-East Baltic speakers (such as the Trzciniec outlier), as we have already discussed in this blog. The unexpected increase in Corded Ware-like ancestry in the Eastern Baltic, coupled with the expected large continuity of hg. R1a-Z283 in the homeland of Balto-Finnic expansions, gives even more support to the known complex system of exogamy along the Baltic coasts, and offers another potential reason for the rise of Baltic-speaking territories in the West Baltic: elite domination.

It is nevertheless important to understand that, even among the most “genetic continuous” regions like Estonia, not a single population in Europe is heir of some ancestral, immutable people. Not in terms of haplogroups, and not in terms of admixture. Balto-Finnic speakers, however continuous they might seem (e.g. in Southern Estonians) aren’t an exception.

After all, this blog was (re)born to fight the currently prevalent sheer stupidity surrounding the simplistic “R1a/steppe ancestry=Indo-European” association, so I wouldn’t like to see it replaced with some other stupid continuity or purity ideas within 10 to 20 years…

Late Uralic stems from East Corded Ware groups

With the currently available tools – linguistics, archaeology, and now genetics -, I don’t think there is any argument to date to question the direct connection of the Late Proto-Uralic expansion with all Eastern Corded Ware groups (i.e. Battle Axe, Fatyanovo-Balanovo, and Abashevo), and thus at least with the unifying A-horizon of Corded Ware and the bottlenecks under R1a-Z645.

NOTE. The only out-group among Corded Ware cultures is the Single Grave culture. It appears to be an early Corded Ware offshoot, reflected in their non-unitary cultural traits (distinct from later unifying waves), in their varied patrilineal clans, and in the short-lasting cultural effect in northern Europe before their complete demise under pressure of expanding Yamna/Bell Beaker peoples from the Danube. The culture’s minimal (if any) effects on succeeding peoples might be seen mostly in the (mainly phonetic) Uralic substrate found in Balto-Slavic – although this may also stem from a more eastern influence, close to the Baltic – and in the contacts of Celtic with Uralic. The huge time depth between this early hypothetic Uralic layer in northern Europe and the emergence of peoples inhabiting these territories in recorded history have no doubt been erroneously interpreted as a lack of Uralic presence in the area.

1) That connection was evident in the Yamna – CWC differences in archaeology, and especially later, with at least Fatyanovo-Balanovo and Abashevo representing the obvious replacement of the Volosovo culture before further expansions of CWC-related groups west and east of the Urals.

The mythical millennia-long continuity of Volosovo hunter-gatherers, including centuries among Corded Ware peoples, as expected lately by the Copenhagen group (and anyone who doesn’t want to question the 1960s association of Indo-European with CWC) must be rejected today in population genomics, as the recent studies of ancient and modern populations show, and as ancient DNA from the region will confirm.

2) In linguistics, the survival of Volosovo as The Uralic-speaking culture was also hardly believable. From Kallio (2015):

While we can say at least something about Uralic substrates in Northeastern Europe, non-Uralic substrates cannot at all easily be identified, because of multiple language shifts, viz. first from non-Uralic to Uralic and then from Uralic to Russian. Yet the Soviet Uralicist Boris Serebrennikov (1956, 1959) argued that there are some non-Uralic substrate toponyms in the Volga-Oka region, but his idea was never taken seriously in the west (cf. Sauvageot 1958), and it pretty soon also sank into oblivion in Russia, even though it can still occasionally pop up there in non-onomastic circles (cf. Napolskikh 1995: 18–19). However, not all the hypotheses on non-Uralic substrates in Northeastern Europe should be rejected (see e.g. Helimski 2001b).

bronze-age-early-languages-east-europe
Tentative map of the distribution of known languages in Eastern Europe during the Early Bronze Age. See full map.

Helimski (2001) argues for a non-Uralic topo-hydronomy in Northern Russia, whose population may have kept their languages up to the Common Era despite the Corded Ware expansion, which is in line with the survival of some non-Indo-European languages everywhere in Europe after the expansion of Yamna and its offshoots:

It should be borne in mind that these [Uralic] hydronyms reached us mainly through Northern Russian and, accordingly, with a tendency to phonetic-morphological adaptation and unification (for river names it is “natural” to be, like the word ‘river’ itself, feminine and to end in -a). Taking into account this circumstance, it may turn out to be non-useless for etymological identification of at least some of the hydronyms on the Finno-Ugric basis.

On the other hand, I wouldn’t exclude the possibility that some parts of this large geographical area were never (completely) Finno-Ugric. The population that created the most important part of the hydronymy of the Russian North could be finally pushed aside or assimilated only at the end of the 1st – beginning of the 2nd millennium AD, during the Russian colonization, retaining the memory of the White-Eyed Chude in its own memory.

NOTE. For more on this non-IE substrate in (especially West) Uralic, see e.g. Zhivlov (2015),

The same non-Uralic substrate is most likely behind most of the shared traits by Mordvinic and Balto-Finnic (see below).

3) In genetics, I don’t think the picture could get any clearer. I don’t know what “Steppe ancestry = Indo-European” proponents expected from 2019, if they expected anything at all (I haven’t seen any coherent model, proposal, or prediction for a long time now), but I doubt the recent results are compatible with any of their implied expectations.

corded-ware-pca-sub-neolithic-europe
Detail of the PCA of the Corded Ware expansion. See full PCA and more related files.

Notice, from the PCA above, how this Baltic Late Neolithic group shows actually a shift from Sredni Stog (see PCA with Sredni Stog) towards typical Khvalynsk-Urals-related ancestry, i.e. populations from eastern European forested regions, derived from hunter-gatherer pottery groups, as I have proposed for a very long time, since the first time a Baltic LN “outlier” appeared. It’s amazing how some amateurs can find 0.1% of any Siberian outlier’s ancestry among Uralians 4,000 years later, but fail to see the direct connection here. The esoteric uses of qpAdm, I guess…

Especially noticeable is the extra WHG-like ancestry and corresponding shift, seen especially marked in late Polish CWC samples, but also in Baltic CWC and especially in one Sweden Battle Axe sample, all of them shifting apparently closer to Pitted Ware and SHG. While that may have been interpreted as an in situ admixture in Scandinavia before, the late Polish CWC samples show likely a resurgence of local populations, so we can assume that both shifts (to SHG- and EHG-like populations) of available CWC samples around the Baltic are clearly part of the WHG:EHG continuum that will be found in the eastern European sub-Neolithic cultures, from Narva to Volosovo.

This WHG-related ancestry is clearly predominant in groups with which Battle Axe peoples admixed, based on the shift towards Pitted Ware, which – I can only guess based on modern Volga Finns – is different from the shift we will see in Netted Ware, more towards the Khvalynsk-Urals cluster. This is in line with the expansion of Battle Axe eastward through coastal areas (West to East Baltic and Finland into Sweden), while Fatyanovo peoples probably emerged from a slightly different route, but also a northern one, if one is to follow archaological similarities and their chronology.

bronze-age-europe-baltic
Detail of the PCA of European Bronze Age populations. See full PCA and more related files.

During the Iron Age, the only peoples that probably shifted strongly (based on modern populations) are West Baltic ones, getting closer to the available Late Trzciniec samples, and even closer to the Trzciniec outlier, i.e. away from the earlier Eastern Corded Ware cluster, and towards Central European groups like Czech EBA or Poland EBA, both of them clearly derived from Bell Beakers, but also admixed with (and thus shifted toward) CW-like populations.

If one looks carefully at the previous PCA on Bronze Age populations, and the next one on Iron Age clusters, it is evident that adding the Swedish LN outlier to East Baltic BA (both strongly related to Battle Axe populations) essentially gives us the continuity of East Baltic BA into the Iron Age. This cluster is continued also in two outliers from Sigtuna, a Viking town close to the Gulf of Finland, known to be an important trading site, 1,500 years later. Not much of a change around the Gulf of Finland, then:

iron-age-eastern-europe
Detail of the PCA of East and North European Iron Age populations. See full PCA and more related files.

Based on the two simplistic Uralic clines one might see described (among the many that certainly existed, from Corded Ware to different Eurasian populations), and just like BOO was for some months fashionable as “Samic”, some may be tempted to say that certain Sintashta or Srubna outliers close to the Urals mark the True Uralic™ peoples. Because, of course they do. Ghost haplogroup N and stuff. And Corded Ware never ever Uralic. Because Gimbutas, and my IE R1a grandfather.

NOTE. Funny thing here: there might be Corded Ware, Iranian, Slavic, Germanic, etc… outliers or out-groups, and they might form the widest genetic clusters ever seen, but they are all of one language, because archaeology and linguistics; however, one “outlier” (also, put your own definition of “outlier” here, let’s say 1% of whatever, and strontium isotope potentially from 100 km away) ca. 600 BC in the Baltic who (surprise!) happens to show hg. N, and he signals the first incoming True Uralic™ speaker from wherever… It won’t be the first or the last time some people resort to “the complexity of Uralic-speaking peoples” in ancestry, just to look for “hg. N = Uralic” like crazy. You only need common sense to understand that this is not how this works. Amateur genomics can’t get more embarrassing than the current “let’s look for ‘Siberian ancestry’ in every individual of haplogroup N” trend. Or maybe it can, and it will, but I can’t see it yet.

If one were to insist on looking for ‘foreign’ contributions among Iron Age Estonians, though, I think one should also check out first archaeology, and then the PC3 (or, more graphically, a 3D plot), to understand what might be happening with the many Uralic clines derived from Corded Ware, before starting to play around with bioinformatic tools to discover a teeny tiny 1% admixture of the wrong population, and rushing to build far-fetched narratives. Apparently, one of the different clines formed roughly between southern (steppe – forest-steppe) and northern (tundra-taiga) populations in Uralians is also seen in some Iron Age Estonian individuals – especially in some late samples from Ingria…This is not my main interest, so I will leave this here for others to keep wasting their time chasing the white whale of the 0.5% of True Uralic™ ancestry in ancient Baltic samples of hg. N.

pca-3d-estonians-iron-age-boo-samic
Still images of the 3D plot of Eurasian samples. Typical PC1 vs. PC2 visualization to the left, and shift of the view to PC3 on the right image. See full PCA and more related files.

An exclusive Volga-Kama homeland for Disintegrating Uralic?

Since I don’t believe in macro-regions of largely continuous ethnolinguistic communities, as I have often said about Slavic (naively associated with prehistoric tribes of Eastern Europe) or Germanic (absurdly considered to be represented by Battle Axe), it is difficult for me to believe that Battle Axe-derived cultures remained of the same Finno-Samic dialects since the Corded Ware expansion…unless we live in Westeros, where everything happens “for thousands of years”.

I have to admit, then, that the now prevalent identification among Uralicists has become quite attractive:

  • Fatyanovo-Balanovo as Finno-Permic:
    • Fatyanovo/Netted Ware with West Uralic (also called Finno-Mordvinic).
    • Balanovo/Chirkovo-Kazan with Central Uralic (Mari-Permic).
  • Abashevo, into the Andronovo-like Horizon through the Seima-Turbino phenomenon, with East Uralic (also Ugro-Samoyedic).

Exactly like the identification of Yamna Hungary – Bell Beaker transition as the North-West Indo-European homeland, it gives us simplicity and small and late ethnolinguistic communities, away from the traditionally overused big and early language territories.

This late homeland would be supported, among others, by:

  • The presence of Indo-Iranian loanwords in Finno-Permic and Ugric (probably also in Samoyedic, either lost, or – much more likely – underresearched), compatible with the immediate contact between Abashevo – Sintashta-Potapovka-Filatovka and Fatyanovo-Balanovo.
  • The supposed expansion of Netted Ware from Fatyanovo to the north-west, which may be explained as the split and expansion of Balto-Finnic and Samic ca. 1900 BC.
  • A longer-lasting Finno-Permic (West+Central Uralic) community contrasting with the early separation of East Uralic.
  • The compatibility of this late expansion with the late expansion of Pre-Germanic from Denmark with the Dagger Period, and of Balto-Slavic with Trzciniec, which puts all three dialects reaching the Baltic Sea in the EBA.

NOTE. I meant to update the linguistic text to include the most recently favoured phylogenetic tree of Uralic languages after Häkkinen (2007, 2009, 2014), which has very quickly become the new normal among Uralicists, but I don’t think I will have enough time to review the necessary papers for that. I am rushing to publish a printed edition, so the text will wind up being a mixture of “traditional” (meaning, basically, pre-2010s) description of Uralic dialects but using modern divisions; say, “West Uralic” instead of “Finno-Samic”. By the way, I am still amazed that none of my reader-haters (or any online user discussing Uralic migrations, for that matter) have come up with the questions that the new division pose, and it supports my suspicion about the complete lack of interest in linguistics of most (a)DNA fans, except for the occasional use of old and free PDFs Googled to support new narratives invented expressly for some qpAdm results…

textile-ceramics-europe-bronze-age
Textile ceramic styles and influence of Bronze Age cultures divided in clusters.

Problems with this Parpola-Carpelan’s (2012-2018) interpretation include:

  • The differentiation between Fennoscandian Textile Ceramics vs. Netted Ware, which is not warranted in archaeology. The assumption that Netted Ware expanded to the Baltic Sea (as Kallio does, following the traditional view) is thus weak, and it was probably a question of cultural contacts coupled with short-distance population movements/exchange in both directions (from the Baltic to the Volga and vice versa). In fact, the culture division relies on some fairly common and technically simple ornamentation patterns, widespread all over northern Europe, even before the Corded Ware expansion, and it is very difficult to separate certain neighboring Textile Ceramics from Netted Ware groups in southern Finland (i.e. Sarsa-Tomitsa groups).
  • The strict and radical direction described for the Netted Ware by Carpelan, as an eastward and northward expansion, within a very short time frame (ca. 1900-1800 BC), based on few radiocarbon dates, which seems to me like a very risky assumption. We know how this kind of descriptions of direction of culture expansion based on radiocarbon dates has turned out in much more complex “packages”, like the Bell Beaker culture… In fact, the earliest dates for Textile Ware are from the East Baltic, earlier than those of Netted Ware.
  • The assumption that Balto-Finnic traits shared with Mordvinic are a) late and b) meaningful for dialectalization of two closely related dialects, when it is clear that both dialects separated quite early. Phonologically Finnic is more conservative, morphologically less so, and the shared traits include a handful of non-Uralic substrate words which can’t be traced to a single common source, hence they were adopted when both languages had already separated… All in all, Finnic – Mordvinic correspondances are not even close to Italo-Celtic ones, which is clearly fully incompatible with a proposal of a Finnic separation from Mordvinic coinciding with the LBA-IA transition.

Especially problematic for Parpola’s model is the lack of genetic impact in Bronze Age or Iron Age Estonians, not reaching a significant level under any possible statistical threshold – which I am sure was quite disappointing for some of my readers -, but is in line with major archaeological continuity of groups the from region, only disturbed in cultural (and Y-chromosome) terms by the expansion of Akozino warrior-traders all over the Baltic Sea. Any proposed population movement will be very difficult to support in genetics, given the Corded Ware-derived populations that we will see in both regions, and the continued Baltic-Volga contacts since the Corded Ware expansion.

Problems with an interpretation of such a small impact in population genomics includes the similarly weak impacts and haplogroup infiltrations that can be seen among populations basically everywhere in Eurasia, during any given period, and much greater genetic impacts that are supposed to be (or that were certainly) followed by ethnolinguistic continuity.

akozino-malar-axes-fennoscandia
Distribution of the Akozino-Mälar axes according to Sergej V. Kuz’minykh (1996: 8, Abb. 2).

The Battle Axe question

From Kallio (2015), about choosing a tentative homeland for Proto-Uralic:

(…) linguistically uniform Proto-Uralic would have been spoken in the Volga-Oka region until the mid-third millennium BC when the Proto-Uralic-speaking area would have expanded to the Volga-Kama region as well. By the end of the same millennium, this expansion would have led to the earliest dialectal splits within Uralic into Finno-Mordvin, Mari-Permic, and Ugro-Samoyed. The splitting up of these three soon followed during the early second millennium BC when the Uralic-speaking area finally stretched from the Baltic Sea in the west to the Altai mountains in the east. Indeed, no matter where Proto-Uralic was spoken, the branching into the nine well-attested subgroups (viz. Finnic, Saami, Mordvin, Mari, Permic, Hungarian, Mansi, Khanty, and Samoyed) must have taken less than a millennium, because their shared phonological and morphosyntactic isoglosses are rather limited (see Salminen 2002). The traditional view that all this branching would have taken several millennia violates everything linguistic typology teaches us about the rate of language change.

The basic problem of this identification of Fatyanovo-Balanovo as West-Central Uralic and Abashevo as East Uralic is the nature of the Battle Axe culture, including the Bronze Age East Baltic and Gulf of Finland area. Even if it is accepted that Fatyanovo-Balanovo represented all Western groups, Battle Axe must have represented West Uralic-like dialects.

The ethnolinguistic identification of Battle Axe depends ultimately on the nature of contacts of Fatyanovo/Netted Ware with Battle Axe/Textile Ceramics. If both groups were close and interacted profusely, as it seems, it doesn’t seem granted that we will be able to distinguish a close Para-West Uralic dialect of Scandinavia from the actual expanding Balto-Finnic and Samic dialects, if they were actually linked to the Netted Ware expansion. Also from Kallio (2015):

No doubt the most convincing substrate theory has recently been put forward by the Saami Uralicist Ante Aikio (2004), who has not only rehabilitated but also improved the old idea of a non-Uralic substrate in Saami. His study shows that there were still non-Uralic languages spoken in Northern Fennoscandia as recently as the first millennium AD. Most of all, they were not only genetically non-Uralic but also typologically non-Uralic-looking, bearing a closer resemblance to the so-called Palaeo-European substrates (for which see e.g. Schrijver 2001; Vennemann 2003).

In comparison, the case of Finnic is much more difficult. The fact that Proto-Uralic was not spoken in the East Baltic region means that this area must have originally been non-Uralic-speaking, but so far the evidence for a non-Uralic substrate in Finnic has consisted of appellatives and proper names with no etymology (cf. Ariste 1971; Saarikivi 2004a). Contrary to the proposed substrate words in Saami, those in Finnic show no structural non-Uralisms, as if they had indeed been borrowed from some genetically related or at least typologically similar languages, as I suggested above. Also none of them is more recent than the Middle Proto-Finnic stage, which makes them at least two millennia old. All this agrees with archaeological evidence discussed earlier that the Uralicization of the East Baltic region occurred during the Bronze Age (ca. 1900–500 BC).

The discussion of the paper continues with an unsuccessful attempt to find a hypothetical ancient Indo-European substrate that Kallio believes must be associated with the expansion of Corded Ware, in line with the traditional belief. For example, the often mentioned – almost folk etymology-like, unsurprisingly popular among amateurs – ‘Neva’ as derived from IE “young” is logically rejected…Unlike Parpola, Kallio’s view seems to be confident that Netted Ware (as Textile Ware) expanded into the East Baltic, on both sides of the Gulf of Finland, already during the Bronze Age.

As it has become apparent in population genomics, none of them was right, and Textile Ceramics will essentially show – like Netted Ware – a large genetic continuity of Corded Ware peoples in the whole north-eastern European forest zone – despite small regional population movements, obviously -, which necessarily implies that the whole Corded Ware culture – and not only Fatyanovo-Balanovo and Abashevo – were Uralic-speaking territories.

The similarities in terms of culture and Y-DNA bottlenecks between Battle Axe and Fatyanovo-Balanovo also imply that the linguistic differences between these groups were probably not many, and became strongly divided only after their territorial division. Continued contacts between Battle Axe- and Fatyanovo-derived groups can explain the proposed contacts (Finnic with Samic, Finnic with Mordvinic) after their linguistic-but-not-physical separation.

east-european-fatyanovocwc
East European movement directions (arrows) of the representatives of the Central European Corded Ware Culture (according to I.I. Artemenko).

Battle Axe spoke “Para-Balto-Finnic”?

The Balto-Finnic-speaking nature of Battle Axe is thus supported by:

  • The lack of non-Uralic substrates in Balto-Finnic territory (Kallio 2015).
  • The early separation of Samic and Finnic from Mordvinic, and the virtual identity of Proto-West-Uralic and Proto-Uralic, which suggests that Proto-Uralic spread fast (Parpola 2012).
  • The scarce non-Uralic topo-hydronymy in the East Baltic and around the Gulf of Finland (Saarikivi 2004), comparable to that on the Upper Volga region.
  • The strong influence of a Balto-Finnic-like substrate on Pre-Germanic (or, in Kallio’s opinion, the same Scandinavian substrate influencing both Germanic and Balto-Finnic at the same time), and the continued influence of Balto-Finnic on Proto-Baltic and Proto-Slavic.
  • The continued influence of Corded Ware-derived groups in central-east Sweden in Finland and the East Baltic in terms of agricultural innovations appearing in the LBA, compatible with Schrijver’s proposal of intermediate Germanic-shifted Balto-Finnic groups and Balto-Finnic groups influenced by their pronunciation.
  • The intense Palaeo-Germanic and late Balto-Slavic / early Proto-Baltic superstrate on Balto-Finnic, which place all three dialects around the Baltic Sea since the Early Bronze Age.
  • The easy replacement of a hypothetic Para-Balto-Finnic dialect by incoming Proto-Balto-Finnic-speaking peoples (say, with textile ceramics), without much linguistic impact.

In fact, the continuous contacts of the East Baltic with the Volga, and especially the close interaction with Akozino warrior-traders just before the Tarand-grave period, could be the actual origin of the recent (if any) Finnic-Mordvinic connections that need to be traced back to the LBA-IA (maybe here the number ‘ten’), since most of them can be related to a Pit-Comb Ware culture substrate and earlier contacts through the forest zone, which Samic (due to its early split and presence to the north of the Gulf of Finland during the BA) does not share. In fact, some of them can be traced back to Balto-Finnic first

These are the most often mentioned, in order of descending relevance for a shared ancient community:

  • Noun paradigms and the form and function of individual cases.
  • The geminate *mm (foreign to Proto-Uralic before the development of Fennic under Germanic influence) and other non-Uralic consonant clusters.
  • The change of numeral *luka ‘ten’ with (non-Uralic) *kümmen.
  • The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature.

It’s not only a question of quantity. Are these shared Mordvinic – Balto-Finnic traits really more relevant than, say, those between Italo-Celtic, which are supposed to have formed a community for a very short period at the end of the 3rd millennium around the Alps? Are these traits even sufficient to propose a common early Mordvinic-Finnic group within West Uralic, rather than loose Mordvinic – Balto-Finnic contacts, i.e. contacts between East Baltic (Textile Ceramics) and Volga-Kama (Netted Ware)?

Based on the alternative (Kallio’s) view of continued contacts between Textile Ceramics groups, even without knowing anything about linguistics, you can guess that Parpola is spinning very thin when assuming that these changes suggest that Balto-Finnic may have expanded with Akozino warrior-traders, separating thus ca. 800 BC from Mordvinic…

Genetic findings now clearly help dismiss any meaningful population impact in the LBA-IA transition, although any linguist can obviously argue for linguistic change in spite of major genetic continuity. But then we are stuck in the pre-ancient DNA era, so what’s ancient DNA for.

netted-ware-textile-ceramics
Middle Bronze Age cultures of Eastern Europe.

Genetic continuity = language continuity?

In the end, it’s very difficult to say how much language continuity there is around Estonia since the arrival of Corded Ware peoples. Looking at Modern Estonians, they have been clearly influenced by recent contacts with Baltic- and Germanic-speaking peoples clustering to the south-west in the PCA. They seem to have also received contacts from north(-east)ern peoples, likely from Finland, evidenced by their shifts toward the modern Estonian cluster during and after the Middle Ages, with a slight increase in Siberian ancestry and N1c subclades associated with Lovozero Ware. How much language change did these contacts bring? Maybe an expansion of Gulf of Finland Finnic (Northern Estonian) over Inland Finnic (Southern Estonian) and Gulf of Riga Finnic (Livonian)? Difficult to know, exactly, but, in the traditional view of Balto-Finnic dialectal distribution among Uralicists like Kallio, possibly no change at all.

So, if the obvious changes in the Estonia_MA cluster relative to Estonia_IA cluster and Estonia_Modern relative to Estonia_MA do not represent radical language change…Why would Estonia_IA represent a change relative to Estonia_BA, when it is statistically basically the same? Or Estonia_BA relative to CWC_Baltic? Because of the infiltration of haplogroup N1c around the whole Baltic? Because of the occasional 1% “Siberian” ancestry in some non-locals of varied haplogroups across the whole Baltic area?

In spite of all this, the amount of special pleading we are seeing among openly Nordicist amateurs when discussing the Uralic homeland relative to the Indo-European question in genetics has become a matter of plain willful ignorance. Like the living corpses of the Anatolian homeland, the Armenian homeland, the OIT proponents, or the nativist Basque R1b association, the personal involvement in the revival of “R1a=Indo-European” and “N=Uralic” trends is just painful to watch.

[Next post in this line, if I manage to make time for it: “Genetic (dis)continuity in Central Europe“. Let’s see if early Balts and early Slavs, as well as Germanic peoples, show a cluster closer to Danubian EBA (viz. Maros), Hungary-Balkans BA, and Urnfield-related samples than their predecessors in their areas, i.e. away from East Corded Ware groups… If you want, you can enjoy for the moment the new PCAs I could get done and the tentative map of languages in the Early Bronze Age, that will probably give you the right idea about early Indo-European and Uralic population movements]

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

Related

Common pitfalls in human genomics and bioinformatics: ADMIXTURE, PCA, and the ‘Yamnaya’ ancestral component

invasion-from-the-steppe-yamnaya

Good timing for the publication of two interesting papers, that a lot of people should read very carefully:

ADMIXTURE

Open access A tutorial on how not to over-interpret STRUCTURE and ADMIXTURE bar plots, by Daniel J. Lawson, Lucy van Dorp & Daniel Falush, Nature Communications (2018).

Interesting excerpts (emphasis mine):

Experienced researchers, particularly those interested in population structure and historical inference, typically present STRUCTURE results alongside other methods that make different modelling assumptions. These include TreeMix, ADMIXTUREGRAPH, fineSTRUCTURE, GLOBETROTTER, f3 and D statistics, amongst many others. These models can be used both to probe whether assumptions of the model are likely to hold and to validate specific features of the results. Each also comes with its own pitfalls and difficulties of interpretation. It is not obvious that any single approach represents a direct replacement as a data summary tool. Here we build more directly on the results of STRUCTURE/ADMIXTURE by developing a new approach, badMIXTURE, to examine which features of the data are poorly fit by the model. Rather than intending to replace more specific or sophisticated analyses, we hope to encourage their use by making the limitations of the initial analysis clearer.

The default interpretation protocol

Most researchers are cautious but literal in their interpretation of STRUCTURE and ADMIXTURE results, as caricatured in Fig. 1, as it is difficult to interpret the results at all without making several of these assumptions. Here we use simulated and real data to illustrate how following this protocol can lead to inference of false histories, and how badMIXTURE can be used to examine model fit and avoid common pitfalls.

admixture-protocol
A protocol for interpreting admixture estimates, based on the assumption that the model underlying the inference is correct. If these assumptions are not validated, there is substantial danger of over-interpretation. The “Core protocol” describes the assumptions that are made by the admixture model itself (Protocol 1, 3, 4), and inference for estimating K (Protocol 2). The “Algorithm input” protocol describes choices that can further bias results, while the “Interpretation” protocol describes assumptions that can be made in interpreting the output that are not directly supported by model inference

Discussion

STRUCTURE and ADMIXTURE are popular because they give the user a broad-brush view of variation in genetic data, while allowing the possibility of zooming down on details about specific individuals or labelled groups. Unfortunately it is rarely the case that sampled data follows a simple history comprising a differentiation phase followed by a mixture phase, as assumed in an ADMIXTURE model and highlighted by case study 1. Naïve inferences based on this model (the Protocol of Fig. 1) can be misleading if sampling strategy or the inferred value of the number of populations K is inappropriate, or if recent bottlenecks or unobserved ancient structure appear in the data. It is therefore useful when interpreting the results obtained from real data to think of STRUCTURE and ADMIXTURE as algorithms that parsimoniously explain variation between individuals rather than as parametric models of divergence and admixture.

For example, if admixture events or genetic drift affect all members of the sample equally, then there is no variation between individuals for the model to explain. Non-African humans have a few percent Neanderthal ancestry, but this is invisible to STRUCTURE or ADMIXTURE since it does not result in differences in ancestry profiles between individuals. The same reasoning helps to explain why for most data sets—even in species such as humans where mixing is commonplace—each of the K populations is inferred by STRUCTURE/ADMIXTURE to have non-admixed representatives in the sample. If every individual in a group is in fact admixed, then (with some exceptions) the model simply shifts the allele frequencies of the inferred ancestral population to reflect the fraction of admixture that is shared by all individuals.

Several methods have been developed to estimate K, but for real data, the assumption that there is a true value is always incorrect; the question rather being whether the model is a good enough approximation to be practically useful. First, there may be close relatives in the sample which violates model assumptions. Second, there might be “isolation by distance”, meaning that there are no discrete populations at all. Third, population structure may be hierarchical, with subtle subdivisions nested within diverged groups. This kind of structure can be hard for the algorithms to detect and can lead to underestimation of K. Fourth, population structure may be fluid between historical epochs, with multiple events and structures leaving signals in the data. Many users examine the results of multiple K simultaneously but this makes interpretation more complex, especially because it makes it easier for users to find support for preconceptions about the data somewhere in the results.

In practice, the best that can be expected is that the algorithms choose the smallest number of ancestral populations that can explain the most salient variation in the data. Unless the demographic history of the sample is particularly simple, the value of K inferred according to any statistically sensible criterion is likely to be smaller than the number of distinct drift events that have practically impacted the sample. The algorithm uses variation in admixture proportions between individuals to approximately mimic the effect of more than K distinct drift events without estimating ancestral populations corresponding to each one. In other words, an admixture model is almost always “wrong” (Assumption 2 of the Core protocol, Fig. 1) and should not be interpreted without examining whether this lack of fit matters for a given question.

admixture-pitfalls
Three scenarios that give indistinguishable ADMIXTURE results. a Simplified schematic of each simulation scenario. b Inferred ADMIXTURE plots at K= 11. c CHROMOPAINTER inferred painting palettes.

Because STRUCTURE/ADMIXTURE accounts for the most salient variation, results are greatly affected by sample size in common with other methods. Specifically, groups that contain fewer samples or have undergone little population-specific drift of their own are likely to be fit as mixes of multiple drifted groups, rather than assigned to their own ancestral population. Indeed, if an ancient sample is put into a data set of modern individuals, the ancient sample is typically represented as an admixture of the modern populations (e.g., ref. 28,29), which can happen even if the individual sample is older than the split date of the modern populations and thus cannot be admixed.

This paper was already available as a preprint in bioRxiv (first published in 2016) and it is incredible that it needed to wait all this time to be published. I found it weird how reviewers focused on the “tone” of the paper. I think it is great to see files from the peer review process published, but we need to know who these reviewers were, to understand their whiny remarks… A lot of geneticists out there need to develop a thick skin, or else we are going to see more and more delays based on a perceived incorrect tone towards the field, which seems a rather subjective reason to force researchers to correct a paper.

PCA of SNP data

Open access Effective principal components analysis of SNP data, by Gauch, Qian, Piepho, Zhou, & Chen, bioRxiv (2018).

Interesting excerpts:

A potential hindrance to our advice to upgrade from PCA graphs to PCA biplots is that the SNPs are often so numerous that they would obscure the Items if both were graphed together. One way to reduce clutter, which is used in several figures in this article, is to present a biplot in two side-by-side panels, one for Items and one for SNPs. Another stratagem is to focus on a manageable subset of SNPs of particular interest and show only them in a biplot in order to avoid obscuring the Items. A later section on causal exploration by current methods mentions several procedures for identifying particularly relevant SNPs.

One of several data transformations is ordinarily applied to SNP data prior to PCA computations, such as centering by SNPs. These transformations make a huge difference in the appearance of PCA graphs or biplots. A SNPs-by-Items data matrix constitutes a two-way factorial design, so analysis of variance (ANOVA) recognizes three sources of variation: SNP main effects, Item main effects, and SNP-by-Item (S×I) interaction effects. Double-Centered PCA (DC-PCA) removes both main effects in order to focus on the remaining S×I interaction effects. The resulting PCs are called interaction principal components (IPCs), and are denoted by IPC1, IPC2, and so on. By way of preview, a later section on PCA variants argues that DC-PCA is best for SNP data. Surprisingly, our literature survey did not encounter even a single analysis identified as DC-PCA.

The axes in PCA graphs or biplots are often scaled to obtain a convenient shape, but actually the axes should have the same scale for many reasons emphasized recently by Malik and Piepho [3]. However, our literature survey found a correct ratio of 1 in only 10% of the articles, a slightly faulty ratio of the larger scale over the shorter scale within 1.1 in 12%, and a substantially faulty ratio above 2 in 16% with the worst cases being ratios of 31 and 44. Especially when the scale along one PCA axis is stretched by a factor of 2 or more relative to the other axis, the relationships among various points or clusters of points are distorted and easily misinterpreted. Also, 7% of the articles failed to show the scale on one or both PCA axes, which leaves readers with an impressionistic graph that cannot be reproduced without effort. The contemporary literature on PCA of SNP data mostly violates the prohibition against stretching axes.

pca-how-to
DC-PCA biplot for oat data. The gradient in the CA-arranged matrix in Fig 13 is shown here for both lines and SNPs by the color scheme red, pink, black, light green, dark green.

The percentage of variation captured by each PC is often included in the axis labels of PCA graphs or biplots. In general this information is worth including, but there are two qualifications. First, these percentages need to be interpreted relative to the size of the data matrix because large datasets can capture a small percentage and yet still be effective. For example, for a large dataset with over 107,000 SNPs for over 6,000 persons, the first two components capture only 0.3693% and 0.117% of the variation, and yet the PCA graph shows clear structure (Fig 1A in [4]). Contrariwise, a PCA graph could capture a large percentage of the total variation, even 50% or more, but that would not guarantee that it will show evident structure in the data. Second, the interpretation of these percentages depends on exactly how the PCA analysis was conducted, as explained in a later section on PCA variants. Readers cannot meaningfully interpret the percentages of variation captured by PCA axes when authors fail to communicate which variant of PCA was used.

Conclusion

Five simple recommendations for effective PCA analysis of SNP data emerge from this investigation.

  1. Use the SNP coding 1 for the rare or minor allele and 0 for the common or major allele.
  2. Use DC-PCA; for any other PCA variant, examine its augmented ANOVA table.
  3. Report which SNP coding and PCA variant were selected, as required by contemporary standards in science for transparency and reproducibility, so that readers can interpret PCA results properly and reproduce PCA analyses reliably.
  4. Produce PCA biplots of both Items and SNPs, rather than merely PCA graphs of only Items, in order to display the joint structure of Items and SNPs and thereby to facilitate causal explanations. Be aware of the arch distortion when interpreting PCA graphs or biplots.
  5. Produce PCA biplots and graphs that have the same scale on every axis.

I read the referenced paper Biplots: Do Not Stretch Them!, by Malik and Piepho (2018), and even though it is not directly applicable to the most commonly available PCA graphs out there, it is a good reminder of the distorting effects of stretching. So for example quite recently in Krause-Kyora et al. (2018), where you can see Corded Ware and BBC samples from Central Europe clustering with samples from Yamna:

NOTE. This is related to a vertical distorsion (i.e. horizontal stretching), but possibly also to the addition of some distant outlier sample/s.

pca-cwc-yamna-bbc
Principal Component Analysis (PCA) of the human Karsdorf and Sorsum samples together with previously published ancient populations projected on 27 modern day West Eurasian populations (not shown) based on a set of 1.23 million SNPs (Mathieson et al., 2015). https://doi.org/10.7554/eLife.36666.006

The so-called ‘Yamnaya’ ancestry

Every time I read papers like these, I remember commenters who kept swearing that genetics was the ultimate science that would solve anthropological problems, where unscientific archaeology and linguistics could not. Well, it seems that, like radiocarbon analysis, these promising developing methods need still a lot of refinement to achieve something meaningful, and that they mean nothing without traditional linguistics and archaeology… But we already knew that.

Also, if this is happening in most peer-reviewed publications, made by professional geneticists, in journals of high impact factor, you can only wonder how many more errors and misinterpretations can be found in the obscure market of so many amateur geneticists out there. Because amateur geneticist is a commonly used misnomer for people who are not geneticists (since they don’t have the most basic education in genetics), and some of them are not even ‘amateurs’ (because they are selling the outputs of bioinformatic tools)… It’s like calling healers ‘amateur doctors’.

NOTE. While everyone involved in population genetics is interested in knowing the truth, and we all have our confirmation (and other kinds of) biases, for those who get paid to tell people what they want to hear, and who have sold lots of wrong interpretations already, the incentives of ‘being right’ – and thus getting involved in crooked and paranoid behaviour regarding different interpretations – are as strong as the money they can win or loose by promoting themselves and selling more ‘product’.

As a reminder of how badly these wrong interpretations of genetic results – and the influence of the so-called ‘amateurs’ – can reflect on research groups, yet another turn of the screw by the Copenhagen group, in the oral presentations at Languages and migrations in pre-historic Europe (7-12 Aug 2018), organized by the Copenhagen University. The common theme seems to be that Bell Beaker and thus R1b-L23 subclades do represent a direct expansion from Yamna now, as opposed to being derived from Corded Ware migrants, as they supported before.

NOTE. Yes, the “Yamna → Corded Ware → Únětice / Bell Beaker” migration model is still commonplace in the Copenhagen workgroup. Yes, in 2018. Guus Kroonen had already admitted they were wrong, and it was already changed in the graphic representation accompanying a recent interview to Willerslev. However, since there is still no official retraction by anyone, it seems that each member has to reject the previous model in their own way, and at their own pace. I don’t think we can expect anyone at this point to accept responsibility for their wrong statements.

So their lead archaeologist, Kristian Kristiansen, in The Indo-Europeanization of Europé (sic):

kristiansen-migrations
Kristiansen’s (2018) map of Indo-European migrations

I love the newly invented arrows of migration from Yamna to the north to distinguish among dialects attributed by them to CWC groups, and the intensive use of materials from Heyd’s publications in the presentation, which means they understand he was right – except for the fact that they are used to support a completely different theory, radically opposed to those defended in Heyd’s model

Now added to the Copenhagen’s unending proposals of language expansions, some pearls from the oral presentation:

  • Corded Ware north of the Carpathians of R1a lineages developed Germanic;
  • R1b borugh [?] Italo-Celtic;
  • the increase in steppe ancestry on north European Bell Beakers mean that they “were a continuation of the Yamnaya/Corded Ware expansion”;
  • Corded Ware groups [] stopped their expansion and took over the Bell Beaker package before migrating to England” [yep, it literally says that];
  • Italo-Celtic expanded to the UK and Iberia with Bell Beakers [I guess that included Lusitanian in Iberia, but not Messapian in Italy; or the opposite; or nothing like that, who knows];
  • 2nd millennium BC Bronze Age Atlantic trade systems expanded Proto-Celtic [yep, trade systems expanded the language]
  • 1st millennium BC expanded Gaulish with La Tène, including a “Gaulish version of Celtic to Ireland/UK” [hmmm, dat British Gaulish indeed].

You know, because, why the hell not? A logical, stable, consequential, no-nonsense approach to Indo-European migrations, as always.

Also, compare still more invented arrows of migrations, from Mikkel Nørtoft’s Introducing the Homeland Timeline Map, going against Kristiansen’s multiple arrows, and even against the own recent fantasy map series in showing Bell Beakers stem from Yamna instead of CWC (or not, you never truly know what arrows actually mean):

corded-ware-migrations
Nørtoft’s (2018) maps of Indo-European migrations.

I really, really loved that perennial arrow of migration from Volosovo, ca. 4000-800 BC (3000+ years, no less!), representing Uralic?, like that, without specifics – which is like saying, “somebody from the eastern forest zone, somehow, at some time, expanded something that was not Indo-European to Finland, and we couldn’t care less, except for the fact that they were certainly not R1a“.

This and Kristiansen’s arrows are the most comical invented migration routes of 2018; and that is saying something, given the dozens of similar maps that people publish in forums and blogs each week.

NOTE. You can read a more reasonable account of how haplogroup R1b-L51 and how R1-Z645 subclades expanded, and which dialects most likely expanded with them.

We don’t know where these scholars of the Danish workgroup stand at this moment, or if they ever had (or intended to have) a common position – beyond their persistent ideas of Yamnaya™ ancestral component = Indo-European and R1a must be Indo-European – , because each new publication changes some essential aspects without expressly stating so, and makes thus everything still messier.

It’s hard to accept that this is a series of presentations made by professional linguists, archaeologists, and geneticists, as stated by the official website, and still harder to imagine that they collaborate within the same professional workgroup, which includes experienced geneticists and academics.

I propose the following video to close future presentations introducing innovative ideas like those above, to help the audience find the appropriate mood:

Related

On the origin and spread of haplogroup R1a-Z645 from eastern Europe

indo-european-uralic-migrations-corded-ware

In my recent post about the origin and expansion of haplogroup R1b-L51, Chetan made an interesting comment on the origin and expansion of R1a-Z645. Since this haplogroup is also relevant for European history and dialectal North-West Indo-European and Indo-Iranian expansion, I feel compelled to do a similar post, although the picture right now is more blurry than that of R1b-L51.

I find it interesting that many geneticists would question the simplistic approach to the Out of Africa model as it is often enunciated, but they would at the same time consider the current simplistic model of Yamna expansion essentially right; a model – if anyone is lost here – based on proportions of the so-called Yamnaya™ ancestral component, as found in a small number of samples, from four or five Eneolithic–Chalcolithic cultures spanning more than a thousand years.

The “75% Yamnaya ancestry of Corded Ware”, which has been given so much publicity since 2015, made geneticists propose a “Yamna → Corded Ware → Únětice / Bell Beaker” migration model, in order of decreasing Yamnaya proportions. Y-DNA and solid archaeological models suggested that this model was wrong, and recent findings have proven it was. In fact, the CWC sample closest to Yamna was a late outlier of Esperstedt in Central Europe, whose ancestry is most likely directly related to Yamna settlers from Hungary.

These wrong interpretations have been now substituted by data from two new early samples from the Baltic, which cluster closely to Yamna, and which – based on the Y-DNA and PCA cluster formed by all Corded Ware samples – are likely the product of female exogamy with Yamna peoples from the neighbouring North Pontic region (as we are seeing, e.g. in the recent Nikitin et al. 2018).

NOTE. There is also another paper from Nikitin et al. (2017), with more ancient mtDNA, “Subdivisions of haplogroups U and C encompass mitochondrial DNA lineages of Eneolithic-Early Bronze Age Kurgan populations of western North Pontic steppe”. Link to paper (behind paywall). Most interesting data is summarized in the following table:

yamna-corded-ware-mtdna

Even after the publication of Olalde et al. (2018) and Wang et al. (2018) – where expanding Yamna settlers and Bell Beakers are clearly seen highly admixed within a few generations, and are found spread across a wide Eurasian cline (sharing one common invariable trait, the paternally inherited haplogroup, as supported by David Reich) – fine-scale studies of population structure and social dynamics is still not a thing for many, even though it receives more and more advocates among geneticists (e.g. Lazaridis, or Veeramah).

NOTE. I have tried to explain, more than once, that the nature and origin of the so-called “Yamnaya ancestry” (then “steppe ancestry”, and now subdivided further as Steppe_EMBA and Steppe_MLBA) is not known with precision before Yamna samples of ca. 3000 BC, and especially that it is not necessarily a marker of Indo-European speakers. Why some people are adamant that steppe ancestry and thus R1a must be Indo-European is mostly related to a combination of grandaddy’s haplogroup, the own modern ethnolinguistic attribution, and an aversion to sharing grandpa with other peoples and cultures.

In the meantime, we are seeing the “Yamnaya proportion” question often reversed: “how do we make Corded Ware stem from Yamna, now that we believed it?”. This is a funny circular reasoning, akin to the one used by proponents of the Franco-Cantabrian origin of R1b, when they look now at EEF proportions in Iberian R1b-L23 samples. It seems too comic to be true.

R1a and steppe ancestry

The most likely origin of haplogroup R1a-Z645 is to be found in eastern Europe. Samples published in the last year support this region as a sort of cradle of R1a expansions:

  • I1819, Y-DNA R1a1-M459, mtDNA U5b2, Ukraine Mesolithic ca. 8825-8561 calBCE, from Vasilievka.
  • I5876, Y-DNA R1a, mtDNA U5a2a, Ukraine Mesolithic 7040-6703 calBCE, from Dereivka.
  • I0061, hg R1a1-M459 (xR1a1a-M17), mtDNA C1, ca. 6773-6000 calBCE (with variable dates), from Yuzhnyy Oleni Ostrov in Karelia.
  • Samples LOK_1980.006 and LOK_1981.024.01, of hg MR1a1a-M17, mtDNA F, Baikalic cultures, dated ca. 5500-5000 BC.
  • Sample I0433, hg R1a1-M459(xM198), mtDNA U5a1i, from Samara Eneolithic, ca. 5200-4000 BCE
  • Samples A3, A8, A9, of hg R1a1-M459, mtDNA H, from sub-Neolithic cultures (Comb Ware and Zhizhitskaya) at Serteyea, although dates (ca. 5th-3rd millennium BC) need possibly a revision (from Chekunova 2014).

NOTE. The fact that Europe is better sampled than North Asia, coupled with the finding of R1a-M17 in Baikalic cultures, poses some problems as to the precise origin of this haplogroup and its subclades. While the first (Palaeolithic or Mesolithic) expansion was almost certainly from Northern Eurasia to the west – due to the Mal’ta sample – , it is still unknown if the different subclades of R1a in Europe are the result of local developments, or rather different east—west migrations through North Eurasia.

Y-Full average estimates pointed to R1a-M417 formation ca. 6500 BC, TMRCA ca. 3500 BC, and R1a-Z645 formation ca. 3300 BC, TMRCA ca. 2900 BC, so the most likely explanation was that R1a-Z645 and its subclades – similar to R1b-L23 subclades, but slightly later) expanded quickly with the expansion of Corded Ware groups.

The presence of steppe ancestry in Ukraine Eneolithic sample I6561, of haplogroup R1a-M417, from Alexandria, dated ca. 4045-3974 calBCE, pointed to the forest steppe area and late Sredni Stog as the most likely territory from where the haplogroup related to the Corded Ware culture expanded.

However, the more recent Y-SNP call showing R1a-Z93 (L657) subclade rendered Y-Full’s (at least formation) estimates too young, so we have to rethink the actual origin of both subclades, R1a-Z93 (formation ca. 2900 BC, TMRCA ca 2700 BC), and R1a-Z283 (formation ca. 2900 BC, TMRCA ca. 2800 BC).

Contrary to what we thought before this, then, it is possible that the expansion of Khvalynsk-Novodanilovka chieftains through the steppes, around the mid-5th millennium BC, had something to do with the expansion of R1a-Z645 to the north, in the forest steppe.

We could think that the finding of Z93 in Alexandria after the expansion of Khvalynsk-Novodanilovka chiefs would make it more likely that R1a-Z645 will be found in the North Pontic area. However, given that Lower Mikhailovka and Kvitjana seem to follow a steppe-related cultural tradition, different to forest steppe cultures (like Dereivka and Alexandria), and that forest steppe cultures show connections to neighbouring northern and western forest regions, the rest of the expanding R1a-Z645 community may not be related directly to the steppe at all.

Adding a hypothetical split and expansion of Z645 subclades to the mid-/late-5th millennium could place the expansion of this haplogroup to the north and west, pushed by expanding Middle PIE-speaking steppe peoples from the east:

distribution-horse-scepters
Schematic depiction of the spread of horse-head scepters in the Middle Eneolithic, representing expanding Khvalynsk-Novodanilovka chieftains. See a full version with notes here.

The Złota culture

I have already written about the Podolia-Volhynia region: about the North Pontic steppe cultures in contact with this area, and about the chaotic period of migrations when Corded Ware seem to have first emerged there among multi-directional and multi-ethnic migrants.

This is what Włodarczak (2017) says about the emergence of Corded Ware with ‘steppe features’ after the previous expansion of such features in Central Europe with Globular Amphorae peoples. He refers here to the Złota culture (appearing ca. 2900-2800 BC) in Lesser Poland, believed to be the (or a) transitional stage between GAC and Corded Ware, before the emergence of the full-fledged “Corded Ware package”.

So far, to the north of the Carpathian Mountains, including Polish lands, no graves indicating their relationship with communities of the steppe zone have been found. On the contrary, the funeral rites always display a local, central European nature. However, individual elements typical of steppe communities do appear, such as the “frog-like” arrangement of the body (Fig. 20), or items associated with Pit Grave milieux (cf. Klochko, Kośko 2009; Włodarczak 2014). A spectacular example of the latter is the pointed-base vessel of Pit Grave culture found at the cemetery in Święte, site 11 near Jarosław (Kośko et al. 2012). These finds constitute a confirmation of the importance of the relationships between communities of Pit Grave culture and Corded Ware culture. They are chronologically diverse, although most of them are dated to 2600-2400 BC – that is, to the “classic” period of Corded Ware culture.

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

However, when discussing the relationships with the steppe communities, Polish lands deserve particular attention since part of the groups inhabiting it belonged to the eastern province of Corded Ware culture (cf. Häusler 2014), which neighboured Pit Grave culture both from the east and south. In addition, there was a tradition of varied relationships with the north Pontic zone, which began to intensify from the second half of the 4th millennium BC (Kośko, Szmyt, 2009; Kośko, Klochko, 2009). These connections are especially readable in Małopolska and Kujawy (Kośko 2014; Włodarczak 2014). The emergence of the community of Globular Amphora culture in the north Pontic zone at the end of the 4th and the beginnings of the 3rd millennium BC (Szmyt 1999) became a harbinger of a cultural closening between the worlds of central Europe and the steppe.

The second important factor taking place at that time was the expansion of the people of Pit Grave culture in a westerly direction, along the Danube thoroughfare. As a result of this, also to the south of the Carpathian Mountains, e.g., along the upper Tisza River, a new “kurgan” cultural system was formed. As one outcome, the areas of central Europe, above all Małopolska, found themselves in the vicinity of areas inhabited by communities characterized by new principles of social organization and a new funeral rite. Around 2800 BC these changes became evident in different regions of Poland, with the most numerous examples being documented in south-eastern Poland and Kujawy. The nature of the funeral rite and the features of the material culture perceptible at that time do not have straight forward analogies in the world of north Pontic communities. In this respect, the “A-horizon” is a phenomenon of local, central European origin. The events preceding the emergence of the said horizon (that is, the expansion of the people of Pit Grave culture into the area north of the arc of the Carpathians) are nowadays completely unidentifiable and remain merely an interesting theoretical matter (cf. e.g., Kośko 2000). Therefore, analysis of the archaeological sources cannot confirm the first archaeogenetic analysis suggesting a bond between the communities of the Pit Grave culture and Corded Ware culture (e.g., Haak et al. 2015).

Artefacts of the “A-horizon”, i.e., shaft-hole axes, amphorae (Fig. 21), beakers, and pots with a plastic wavy strip (Fig. 7) are found in different funerary and settlement contexts, sometimes jointly with finds having characteristics of various cultures (e.g., in graves of Złota culture, or at settlements of Rzucewo culture). Hence, they primarily represent a chronological phase (c. 2800-2600 BC), one obviously related to the expansion of a new ideology.

Eastern CWC expansion

Before continuing tracing the Corded Ware culture’s main features, it is worth it to trace first their movement forward in time, as Corded Ware settlers, from Poland to the east.

Circum-Baltic CWC

According to Klochko and Kośko (1998):

The colonizing Neolithic waves are continued by the Circum-Baltic Corded Ware culture, closely related to the traditions of the Single Grave culture and traditions of the Northern European Lowlands. After ca. 2900 BC, certain cultural systems with ‘corded’ traits –genetically related to the catchment area of the south-western Baltic – appear in the drainages of the Nemen, Dvina, Upper Dnieper, and even the Volga. These communities are considered the vector of Neolithisation in the Forest Zone.

east-european-fatyanovocwc
East European movement directions (arrows) of the representatives of the Central European Corded Ware Culture. Modified from I.I. Artemenko.

The picture in the Baltic (Pamariu / Rzucewo) and Finland (Battle Axe) is thus more or less clearly connected with early dates ca. 2900-2800 BC:

There is a clear interaction sphere between the eastern Gulf of Finland area – reaching from Estonia to the areas of present-day Finland and the Karelian Isthmus in Russia –, evidenced e.g. by the sharp-butted axes, derived from the Estonian Karlova axe.

Interesting in this regard is the expansion of the Corded Ware culture in Finland, into a far greater territory than previously thought, that is poorly represented in most maps depicting the extent of the culture in Europe. Here is summary of CWC findings in Finland, using images from Nordqvist and Häkäla (2014):

finland-cwc
Corded Ware culture remains in Finland, excluding the so-called ‘imitations’. [Notice in the top left image the often depicted border of the culture]. Combination of maps from Nordqvist & Häkälä (2014)

Middle Dnieper and Fatyanovo

The earliest Middle Dnieper remains are related to CWC graves between the Upper Vistula and the Bug, containing pottery with Middle Dnieper traits, dated probably ca. 2700 BC, which links it with the expansion of the A-horizon. In fact, during the period ca. 2800-2400 BC, the area of Lesser Poland (with its numerous kurgans and catacomb burials) is considered the western fringe of an area spreading to the east, to the middle Dniester and middle Dnieper river basins, i.e. regions bordering the steppe oecumene. This ‘eastern connection’ of funeral ritual, raw materials, and stylistic traits of artefacts is also identified in some graves of the Polish Lowlands (Włodarczak 2017).

gac-cwc-baltic-dnieper
Cultural situation in Eastern Europe in approximately the middle of the III mill. BC. Key: 1 – areas settled by Globular Amphora culture populations; 2 – areas penetrated by Globular Amphora culture populations; 3 – border between central and eastern group; 4 – Pamariu/Rzucewo culture area; 5 – zone of Pamariu/Rzucewo culture influences; 6 – directions of Comb Pottery culture influence; 7 – Zhizhitskaya culture; 8 – eastern border of “pure” Corded Ware site; 9 – North Belarussian culture; 10 – Middle Dnieper culture; 11 – Fatyanovo culture; 12 – Yamnaya culture; 13 – eastern border of Dniester group; 14 – Kemi-Oba culture and influences; 15 – Foltesti culture; 16 – syncretic sites with evidence of Globular Amphora culture traits (1 – Nida; 2 – Butinge; 3 – Palanga; 4 – Juodkrante; 5 – Azyarnoye; 6 – Mali Rogi; 7 – Prorva; 8 – Strumen/Losha; 9 – Syabrovichi; 10 – Luchin-Zavale; 11 – Lunevo (?); 12 – Belynets; 13 – Losiatyn; 14 – Corpaci; 15 – Ocnita; 16-17 – Camenca; 18 – Marculesti; 19 – Orhei; 20 – Efimovka; 21 – Tatarbunary; 22 – Novoselitsa; 23 – Primorskoye; 24 – Sanzhiyka; 25 – Akkermen; 26 – Maydanetskoye; 27 – Grigorevka; 28 – Kholmskoye; 29 – Purcari; 30 – Roscani; 31 – Semenovka; 32 – Grishevka; 33 – Durna Skela; 34 – Iskovshchina; 35 – Primorskoye); 17 – borders of ecological zones. From Szmyt (2010)

The Fatyanovo (or Fatyanovo-Balanovo) culture was the easternmost group of the Corded Ware culture, and occupied the centre of the Russian Plain, from Lake Ilmen and the Upper Dnieper drainage to the Wiatka River and the middle course of the Volga. From the few available dates, the oldest ones from the plains of the Moskva river, and from the late Volosovo culture containing also Fatyanovo materials, and in combination they show a date of ca. 2700 BC for its appearance in the region. The Volosovo culture of foragers eventually disappeared when the Fatyanovo culture expanded into the Upper and Middle Volga basin.

The origin of the culture is complicated, because it involves at its earliest stage different Corded Ware influences in neighbouring sites, at least on the Moskva river plains (Krenke et al. 2013): some materials (possibly earlier) show Circum-Baltic and Polish features; other sites show a connection to western materials, in turn a bridge to the Middle Dnieper culture. This suggests that groups belonging to different groups of the corded ware tradition penetrated the Moscow region.

The split of subclades Z93 – Z283

If we take into account that the split between R1-Z93 and R1a-Z283 must have happened during the 5th millennium BC, we have R1a-Z93 likely around the middle Dnieper area (as supported by the Alexandria sample), and R1a-Z283 possibly to the north(-west), so that it could have expanded easily into Central Europe, and – through the northern, Baltic region – to the east.

Where exactly lies the division is unclear, but for the moment all reported Circum-Baltic samples with Z645 subclades seem to belong to Z282, while R1a samples from Sintashta/Potapovka (including the Poltavka outlier) point to Abashevo being dominated by R1a-Z93 subclades.

We have to assume, then, that an original east-west split betwen R1a-Z283 and R1a-Z93 turned, in the eastern migrations, into a north-south split between Z282 and Z93, where Finland and Battle Axe in general is going to show Z282, and Middle Dnieper – Abashevo Z93 subclades.

copper-age-early-cwc
Early Copper Age migrations ca. 3100-2600 BC.

I can think of two reasons why this is important:

  1. Depending on how Proto-Corded Ware peoples expanded, we may be talking about one community overcoming the other and imposing its language. Because either
    • clans of both Z93 and Z283 were quite close and kept intense cultural contacts around Dnieper-Dniester area; or
    • if the split is as early as the 5th millennium BC, and both communities separated then without contact, we are probably going to see a difference in the language spoken by both of them.
  2. In any case, the main north-south division of eastern Corded Ware groups is pointing to an important linguistic division within the Uralic-speaking communities, specifically between a Pre-Finno-Ugric and a Pre-Samoyedic one, and potentially between Pre-Finno-Permic and Pre-Ugric.

These may seem irrelevant questions – especially for people interested only in Indo-European migrations. However, for those interested in the history of Eurasian peoples and languages as a whole, they are relevant: even those who support an ‘eastern’ origin of Proto-Uralic, like Häkkinen, or Parpola (who are, by the way, in the minority, because most Uralicists would point to eastern Europe well before the Yamna expansion), place the Finno-Ugric expansion with the Netted Ware culture as the latest possible Finno-Ugric immigrants in Fennoscandia.

The Netted Ware culture

The image below shows the approximate expansion of Corded Ware peoples of Battle Axe traditions in Finland, as well as neighbouring Fennoscandian territories, from ca. 2800 BC until the end of the 3rd millennium. A controversial 2nd (late) wave of the so-called Estonian Corded Ware is popular in texts about this region, but has not been substantiated, and it seems to be a regional development, rather than the product of migrations.

finland-corded-ware
Left: Corded Ware remains in Finland from ca. 2800 BC, according to Nordqvist & Häkälä (2014), combined in a single image. Right: Distribution of the Corded Ware culture within Finland. Mapped (black dots) are finds of typical stone battle axes, used as a proxy (data from [8]). The red isolines indicate average permanent snow cover period from 1981 to 2010 (data from [9]). A recent study estimates the snow cover period ca 4500 years ago would have been 40–50 days less than today [10]. Overlying coloration refers to the lactose persistance (LP) allele gradient in modern northeastern Europe (see the electronic supplementary material, appendix B: Material and methods and table 1, for details); lozenge dots specify the dataset mean points for the triangulation. From Cramp et al. (2014).

As we have seen, Fatyanovo represents the most likely cultural border zone between Circum-Baltic peoples reaching from the Russian Battle Axe to the south, and Middle Dnieper peoples reaching from Abashevo to the north. In that sense, it also represents the most likely border culture between north-western (mainly R1a-Z282) and south-eastern (R1a-Z93) subclades.

With worsening climatic conditions (cooler seasons) at the end of the 3rd millennium, less settlements are apparent in the archaeological record in Finland. After ca. 2000 BC, two CWC-related cultures remain: in the coast, the Kiukainen culture, derived from the original Circum-Baltic Corded Ware settlers, reverts to a subsistence economy which includes hunting and fishing, and keeps mainly settlements (from the best territories) along the coast. In the inland, Netted Ware immigrants eventually appear from the south.

cultures-western-finland
Image modified from Cramp et al. (2014) “The timeline shows the archaeological cultures
discussed here alongside actual sherds sampled and typical vessel forms (after [26–28]) (latter not shown to scale). Distribution maps show the geographical range of (f) Typical Comb Ware, (g) Corded Ware, (h) Kiukainen Ware and (i) Bronze Age cultures in the region (after [10,20,29]).”

The Netted Ware culture emerged in the Upper Volga–Oka region, derived from the Abashevo culture and its interaction with the Seima-Turbino network, and spread ca. 1900-1800 BC to the north into Finland, spreading into eastern regions previously occupied by cultures producing asbestos and organic-tempered wares (Parpola 2018).

NOTE. Those ‘contaminated’ by the Copenhagen fantasy map series may think that Volosovo hunter-gatherers somehow survived the expansion of Fatyanovo-Balanovo and Abashevo, hidden for hundreds of years in the forest, and then reappeared and expanded the Netted Ware culture. Well, they didn’t. At least not in archaeological terms, and certainly not with the genetic data we have.

If we combine all this information, and we think about these peoples in terms of Pre-Finno-Permic and Pre-Ugric languages developing side by side, we get a really interesting picture (see here for Proto-Fennic estimates):

  • The Battle Axe around the Baltic Sea – including the Gulf of Finland and Scandinavia – would be the area of expansion of Pre-Finno-Permic peoples, of R1a-Z283 subclades, which became later concentrated mainly on coastal regions;
  • the southern areas may correspond to Pre-Ugric peoples, which expanded later to the north with Netted Ware (see image below) – their precise subclades may be dependent on what will be found in Fatyanovo;
  • and Pre-Samoyedic peoples (of R1a-Z93 subclades) would have become isolated somewhere in the Cis- or (more likely) Trans-Urals region after 2000 BC, possibly from the interaction of the latest Balanovo stages and the Seima-Turbino phenomenon.
netted-ware-parpola
Distribution of the Netted Ware according to Carpelan (2002: 198). A: Emergence of the Netted Ware on the Upper Volga c. 1900 calBC. B: Spread of Netted Ware by c. 1800 calBC. C: Early Iron Age spread of Netted Ware. (After Carpelan 2002: 198 > Parpola 2012a: 151.)

These communities in contact would have allowed for:

  • the known Indo-Iranian loanwords in Finno-Ugric to spread through a continuum of early dialects formed by Abashevo – Fatyanovo – Battle Axe groups;
  • the Finno-Saamic substrate of Germanic to be associated with Battle Axe groups in Scandinavia;
  • the important Palaeo-Germanic loanwords in Finno-Saamic spreading with long-term contacts (from Pre-Germanic to the Proto-Germanic, and later North Germanic period) through the Baltic Sea, between Scandinavia and the Gulf of Finland;
  • and Tocharian contacts with Samoyedic (although limited, and in part controversial), which point to its early expansion to the east of the Ural Mountains.

On the other hand, if one is inclined to believe that R1a and steppe ancestry do represent Indo-European speakers… which language was spoken from the Gulf of Finland well into the north, the inland, and Karelia, and in Northern Russia, by Corded Ware peoples and their cultural heirs (like Kiukainen or Netted Ware) for almost three thousand years?

Because we know that no other peoples of different haplogroups dominated over eastern Fennoscandia until the Iron Age, and N1c and Siberian ancestry expanded separately, and probably due to late bottlenecks, especially with Fennic peoples expanding recently to the north at the expense of the Saami population.

After the expansion of Bell Beaker peoples, the geographic distribution of late Corded Ware groups in the second half of the 3rd millennium, just before their demise – and before the expansion of Netted Ware to the north – , can be depicted thus as follows:

bronze_age_early_cut
Early Bronze Age Europe.

Territories in cyan must then represent, for some people who believe in an archaic Indo-Slavonic of sorts, the famous Fennoscandian Balto-Slavic to the north (before they were displaced by incoming Finno-Saamic peoples of hg N1c during the Iron Age and up to the Middle Ages); and the also famous Tundra-Forest Indo-Iranian in the Upper Volga area, a great environment for the development of the two-wheeled chariot…

But let’s leave the discussion on imaginary IE dialects for another post, and continue with the real question at hand.

A steppe funerary connection?

Back to Złota as a transitional culture, we have already seen how the corded ware vessels characteristic of the Classic CWC are related to Globular Amphora tradition, and show no break with this culture. It is usually believed that the funerary rites were adopted from steppe influence, too. That is probably right; but it does not mean that it came from Yamna or other coeval (or previous) steppe culture; at least not directly.

NOTE. A similar problem is seen when we read that Mierzanowice or Trzciniec show “Corded Ware” traits from a neighbouring CWC group, when CWC groups disappeared long before these cultures emerged. For cultural groups that are separated centuries from each other, an assertion as to their relationship needs specifics in terms of dates and material connection, or it is plainly wrong.

These are the funerary ritual features from Złota (later specialized in Corded Ware), as described by Włodarczak (2017):

  • Single burial graves; along with the habit of interring the deceased in multiple burial graves, but emphasizing their individual character by careful deposition of the body and personal nature of the grave goods.
  • Grave goods with materials and stylistiscs belonging to an older system (e.g. amber products); and others correlated to the ‘new world’ of the CWC, such as flint products made of the raw materials tipical of Lesser Poland’s CWC, copper ornaments, stone shaft-hole axes, bone and shell ornaments, and characteristic forms of vessels like beakers and amphoras.
  • Military goods, which would become prevalent in later periods, are present in a moderate number, compatible with their lesser importance.
  • There are also cases of the characteristic catacomb (“niche”) graves – with an entrance pit, a more extensive niche, and a narrow corridor leading to a vault – , as well as some individual cases of application of ochre and deformation of skulls.
catacomb-grave-ksiaznice
Catacomb grave no. 2a/06 from Książnice, Złota culture (acc. to Wilk 2013). Image from Włodarczak (2017)

It seems that the Złota funerary tradition was also “transitional”, like corded ware vessels, into the classical Corded Ware ideology. But “transitional” from what exactly? Yamna? Probably not.

The Lublin-Volhynia culture

One needs not look for a too distant culture to find similarities. Włodarczak (2017) points to CWC in south-eastern Poland and Kuyavia showing, by the time of the Yamna expansion, a funeral rite and features of the material culture without straightforward analogies in the world of north Pontic communities, and thus suggests that the “A-horizon” is a local phenomenon of central European origin.

This assertion is interesting, in so far as most Corded Ware samples investigated to date seem to come precisely from an East-Central territory near the Ukraine forest steppe, with a cluster already established by the end of the 5th millennium:

PCA-caucasus-lola-ane-chg
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. An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them.See the original file here.

The following text is from Stanisław Wilk (2018), about the Lublin-Volhynian (and related) cemeteries at Wyciąże and Książnice:

lublin-volhynia-culture
A reach of the Wyciąże-Złotniki group and Lublin-Volhynian culture in the south-eastern Poland and western Ukraine: 1. Area of the Wyciąże-Złotniki group; 2. Area of the Lublin-Volhynian culture. A. Cemetery of the Lublin-Volhynian culture at site 2 in Książnice; B. Cemetery of the Wyciąże-Złotniki group at site 5 in Kraków Nowa Huta-Wyciąże (drawing by S. Wilk based on Zakościelna 2006 and Nowak 2014, on a background downloaded from https://maps-for-free.com/).

Regardless of the differences between the two necropolises (such as the number of burials, the area which has been explored, the orientation and layout of burials), it seems that they have several key elements in common:

  • concentration of graves in separate cemeteries;
  • differentiation of burials with regard to sex (the principle of the ‘left ̶ right’ side, different burial goods for males and females);
  • stratification of graves with regard to the richness of their inventories (this mainly applied to copper artefacts);
  • occurrence of indicators of the richest male burials (a copper dagger in Wyciąże, a copper battle axe, a small axe and a chisel in Książnice);
  • allocation of a separate area for elite burials (the eastern burial area in Książnice, and the southeastern and north-central part of the necropolis in Wyciąże), as well as one for egalitarian burials (the western area in Książnice, and the south-central and western part of the cemetery in Wyciąże).
lublin-volhynian-eneolithic-cemetery
Plan of the Lublin-Volhynian culture cemetery at site 2 in Książnice: 1. female graves; 2. man graves; 3. copper traces; 4. cenothap; 5. cremation grave; 6. partial grave; 7. estimated area of the L-VC cemetery; 8. estimated area of an elite and poor burial fields; 9. area of burials containing copper artefacts (drawing by S. Wilk).

The above-mentioned characteristics prove that the patterns of social and religious behaviours from areas lying beyond the Carpathian Mountains exerted a strong influence on the two societies living in Lesser Poland.

Anna Zakościelna, while describing the similarities between the burial ritual of the late Polgár groups and cultures from areas on the Tisza river and the Lublin-Volhynia culture, claimed that:

a characteristic feature of the burial ritual of both cultures was practicing various group norms, which required different treatment of the deceased depending on their sex, age and social rank. As in the Lublin-Volhynia culture, the opposition ‘male – female’ can the most clearly be observed ̶ particularly, in the consistent positioning of males on the right, and females, on the left side. And, there is much indication that this ritual norm divided the deceased from early childhood (Sofaer Derevensky 1997: 877, Tab. 1; Lichter 2001: 276- 280, 322-323) (Zakościelna 2010: 227-228).

It seems that these observations can also be extended to the Wyciąże-Złotniki group.

Another question is whether the evidence of the influences of the copper civilization observed in both cemeteries emerged as a result of the literal copying of patterns from the south, or whether the latter were only a source of inspiration for local solutions.

Looking at this problem form the perspective of the details of burial ritual, between the Carpathian Basin and Lesser Poland, we can observe clear differences, among others, in the size of cemeteries and orientation of burials. While, in the Carpathian Basin there were large necropolises, consisting of several dozen burials located in rows, with the dominant orientation along the SE-NW and E-W axis (Lichter 2001: Abb. 123, 143; Kadrow 2008: 87); in Lesser Poland there were small cemeteries of several to a dozen or so burials, mostly oriented along the S-N axis (in the Lublin-Volhynia culture; Zakościelna 2010: 66), as well as S-E and NE-SW (in the Wyciąże-Złotniki group; Kaczanowska 2009: 77). Similarly, there are differences in the details of the burial goods. North of the Carpathians, there is a much smaller frequency of copper artefacts, particularly in the group of prestigious, heavy items (battle axes, axes and daggers), as well as a complete lack of objects made of gold. Want is more, the pottery found in the graves has a distinct local character, only supplemented by imitating or imports from areas beyond the Carpathians (Zakościelna 2006: 85; Nowak 2014: 273; a different opinion Kozłowski 2006: 57). Therefore, the suggestion made by Nowak seems right ̶ namely, that these influences were not caused by migrations of groups of the population living on the Tisza river to Lesser Poland, but were rather due to processes of selective cultural transmission (Nowak 2014: 273).

Therefore, the sharing of a similar funerary rite (as happened later between Lublin-Volhynia and Złota), although it shows a strong cultural connection with autochthonous cultures, is obviously not the same as sharing ancestors; and even if it were so, they would not need to be paternal ancestors. But it shows that important Corded Ware cultural traits are local developments, and it disconnects thus still more supposed CWC ‘steppe traits’ from steppe cultures, and connects them with the first steppe-related cultural wave that reached central Europe in the 5th millennium BC.

Prehistoric Pontic—Caspian links

How would a Lublin-Volhynia culture be related to the North Pontic area ca. 4500-3000 BC? We can enjoy the map series of Baltic—Pontic migrations by Viktor Klochko (2009), and make a wild guess:

baltic-pontic-routes
Pontic—Baltic routes of migrations during the Eneolithic. Top left: Linear Pottery expansion. Top right: Funnel Beaker expansion. Bottom left: late Trypillia expansion. Bottom right: GAC expansion.

And then read the account of Sławomir Kadrow, in Exchange of People, Ideas and Things between Cucuteni-Trypillian Complex and Areas of South-Eastern Poland (2016):

In the second half of the 5th millennium BC (horizon 1), communities of the Tripolye culture, phases BI-BII, had contacts with the population of the late (IIa) phase of the Malice culture. The areas settled by both cultural complexes were located at a great distance from each other. The communities of the Tripolye culture adopted selected features of Malice ceramic production (fig 2). This seems to have resulted from marital exchange: on a moderate scale, Tripolye men sought out their wives in the area of the Malice culture and, according to patrilocal marriage customs, the women then moved to the Tripolye settlements, sporadically transferring ready-made ceramic products, so-called imports, to the Tripolye culture. Thus, the wives were responsible for the considerably more numerous imitations of the Malice ceramics and the long-lasting, though selective, traditions of Malice pottery passed down in their new environment. The patrilocal marriage customs involving the Malice women and the Tripolye men (never the other way round), and the fact that pottery was women’s domain, led to the unidirectional transfer of vessels, technology and norms of ceramic production from the Malice culture to the Tripolye culture.

The turn of the 5th and the 4th millennia and the early 4th millennium BC (horizon 2) witnessed the deepening interaction between the populations of the youngest (IIb) phase of the Malice culture and the classic (II) phase of the Lublin-Volhynia culture on the one hand and the communities of phase BII of the Tripolye culture on the other. The Danube and the Tripolye settlement complexes came into contact on the upper Dniester and between the Styr and the Horyn rivers in Volhynia. This helped to continue the previous forms of marital exchange, which resulted in further popularisation of the ceramics and the traditions of ceramic production typical of the Danube cultures, i.e. the Malice and the Lublin-Volhynia cultures, and also the Polgár culture, in the areas settled by the Tripolye cultural complex.

As the civilizational norms of the Eneolithic (Copper) Age became widespread in that period, the forms of interaction described above acquired new elements. The deepening internal diversification of the early Eneolithic communities of the Lublin-Volhynia culture led to a growing demand for prestige objects, which was met with import or imitation of copper artefacts, mainly those from the Carpathian Basin, and with flint tools produced from long blades. That type of flint production depended largely on new technologies derived from the Tripolye culture, as proven by such borrowings as troughlike retouch or the very idea and technology for the production of long flint blades in the Lublin-Volhynia culture. It seems that the influx of Tripolye settlers into flintbearing areas in Volhynia and on the upper Dniester, adjacent to the settlement centres of the late phase of the Malice culture and the Lublin-Volhynia culture, created sufficient conditions for the expanding influence of the Tripolye flint working on the communities of the Eneolithic Lublin-Volhynia culture.

In the mid-4th millennium BC (horizon 3), those forms of interaction between the Danube communities (the late phase of the Lublin-Volhynian culture) and the Tripolye communities (phase CI)were continued. Elements of the Danube pottery still grew in popularity in the Tripolye population, while selected features of the Tripolye flint working were adopted by the Lublin-Volhynia culture.

In that period, the population of the Funnel Beaker culture of the pre-classic and early classic phases (the beginnings of Gródek 1 and Bronocice III), until then absent from those areas, quite quickly drove out and replaced the Danube population in western Volhynia and the upper Dniester basin. This caused significant changes in the forms and intensity of the intercultural interaction, which became fully apparent already in the 2nd half of the 4th millennium BC.

In the following period (horizon 4), the population of the classic phase of the Funnel Beaker culture (Gródek 1, Bronocice III) settled more and more intensively the upper Dniester basin, up to the Hnyla Lypa river, and western Volhynia, up to the Styr river. East of those rivers, the Funnel Beaker settlers created considerable areas where they mixed with settlers from early phase CII of the Tripolye culture. Their coexistence, lasting there for many generations, resulted in deepening the interactions between members of both cultural complexes and in developing entirely new forms of relationships.

(…)

The intensifying interaction between the communities of the Funnel Beaker culture and the Tripolye culture, early phase CII, in the 2nd half of the 4th millennium BC (horizon 4) was an introduction to, and perhaps a condition for, even more frequent contacts in the next period, the first centuries of the 3rd millennium BC (horizon 5). In that case, the interaction was mainly triggered by multidirectional migrations of larger human groups, involving a significant part of the population of all cultures from the areas discussed here. The Tripolye communities of younger phase CII settled Volhynia, its eastern areas in particular, from the south and the south-east, while groups representing the younger phases of the Funnel Beaker culture (Gródek 2), often with Baden features (Bronocice IV and V), moved increasingly into the western part of that region. The Yamna communities expanded along the lower and central Danube to the west, whereas the populations of the late phase of the Baden culture took the opposite direction, reaching as far as Kiev in the northeast, and contributed to the cultural character of the Sofievka group.

The communities of the Globular Amphora culture migrated from the north-west, from eastern Poland, towards the Danube Delta and as far as the Dnieper in the east, while the multicultural population from the areas around the mouth of the Danube moved in the opposite direction, carrying with them cultural elements from Thrace, or even from Anatolia. Some of them returned to the starting point (to south-eastern Poland), bringing with them a new form of pottery, so-called Thuringian amphora, borrowed from the late Trypillian Usatovo group. This resulted in origins of the Złota culture, a cultural phenomenon that gave beginnings to the oldest Corded Ware culture. Inventories of both cultures contained the already mentioned Thuringian amphorae.

lublin-volhynia-alexandria
Graves and cemeteries with gender differentiated burial rites in Europe; A — Hamangia and Varna cultures; B — Tiszapolgar and Bodrogkeresztur cultures; C — Lublin-Volhynia culture; D — Brześć Kujawski culture. Added star symbol with approximate location of the Alexandria site. Modified image from Sławomir Kadrow (2016)

Here is a more recent assessment (2017) of the latest radiocarbon analyses of the available settlements of cultures in the area, published by Marek Novak (announced in a previous post), which gives the following data on Wyciąże-Złotniki, Lublin-Volhynia, and Wyciąże/Niedźwiedź:

This scheme unambiguously suggests both the overlapping and contiguous nature of cultural development in western Lesser Poland within the Middle Neolithic. The basic elements of this development are: 1) the Wyciąże-Złotniki group and the Lublin-Volhynian culture, until c. 3650–3550 cal BC; 2) the Funnel Beaker culture proper, which appeared c. 3750–3700c al BC, and existed until c. 3300–3250 cal BC, perhaps accompanied by the Wyciąże/Niedźwiedź materials from c. 3650–3550 cal BC; and 3) the Baden culture and the Funnel Beaker/Baden assemblages from 3100 and 3300–3100 cal BC, respectively, until 2850–2750 and 2850 cal BC, with – possibly – later Funnel Beaker culture and Wyciąże/ Niedźwiedź materials, existing until c. 3100 cal BC.

The final scheme shows that the Lublin-Volhynian culture could have coincided with the Wyciąże-Złotniki group. In view of the territorial relationship between them, relations from the point of view of material culture, primarily in the field of pottery, become particularly interesting. It is relatively easy to see clear similarities between these units. However, the most evident similarities apply only to some categories of ceramics, including, for example, vessels with Scheibenhenkel handles. What is more, in the period between the late 38th and early 36th centuries BC, the early Funnel Beaker and possibly early Baden influences are superimposed on this Lublin-Volhynian/Wyciąże-Złotniki ‘mix’.

[About Corded Ware: The] development of this unit in central Europe, including western Lesser Poland, [] usually point to c. 2800 cal BC (Włodarczak 2006a). (…) the calibration curve makes it possible to alternatively refer several dates earlier than c. 3100 to c. 2850–2800 cal BC.

Conclusion

There is no direct archaeological link of Lublin-Volhynia-related groups with Corded Ware, beyond the fact that they shared homeland and Central European (‘steppe-related’) traits, as found in the Złota culture. But there is no direct link of Yamna with Corded Ware, either, whether in terms of culture or population.

So, given the evident link of R1a-Z93 and steppe ancestry with the forest steppe ca. 4000 BC, the surrounding North Pontic areas in contact along the Dniester, Dnieper, Bug, and Prut are the best candidates for the appearance of R1a-Z283: steppe cultures to the south and south-west; sub-Neolithic (Comb Ware) groups to the north in the forest zone; and Eneolithic groups to the west and north-west.

Seeing how ‘ancestral components’ and PCA cluster can change within a few generations, the question of the spread of R1a-Z645 subclades is still not settled by a single sample in Alexandria. However, based on the explosive expansions we are seeing from small territories, it would not be surprising to find R1a-Z93 and R1a-Z283 side by side in the same small area within the forest steppe.

NOTE. An archaeological link may not mean anything relevant in genetics, especially – as in this case – when no clear migration event has been traced to date. We have seen exactly that with Kristiansen’s proposal of a long-term genetic admixture of Yamna with Trypillia and GAC to form Corded Ware, which didn’t happen. The cultural and ideological connection of CWC peoples with Lublin-Volhynian tradition may be similar to the already known connection with GAC, and not mean anything in genetic finds; at least in terms of Y-DNA haplogroup.

We believed in the 2000s that Corded Ware represented the expansion of Late Proto-Indo-European, because the modern map of haplogroup R1a showed a distribution similar to how we thought the European and Indo-Iranian languages could have expanded. This has been proven wrong, and that’s what ancient DNA is for; not to confirm the own ideas or models, or to support modern ideologies.

It is impossible to know if R1a-Z645 comes from the steppe, forest steppe, or forest zone, until more samples are published. I don’t think there will be any big surprise, no matter where it is eventually found. By now, adding linguistic reconstruction to archaeological traits, and to the genetic data from Yamna and Corded Ware settlers, the only clear pattern is that patrilineal clans expanded, during the Final Eneolithic / Chalcolithic:

  • Late Proto-Indo-European with Yamna and R1b-L23 subclades, given the known genomic data from Khvalynsk, Yamna, Afanasevo, Bell Beaker, Catacomb, and Poltavka—Sintashta/Potapovka.
  • Uralic with Corded Ware and R1a-Z645 subclades, given the known genomic data from Fennoscandia and the Forest Zone.

Everything else is just wishful thinking at this moment.

See also

Related

Minimal Corded Ware culture impact in Scandinavia – Bell Beakers the unifying maritime elite

copper-age-late-bell-beaker

Chapter The Sea and Bronze Age Transformations, by Christopher Prescott, Anette Sand-Eriksen, and Knut Ivar Austvoll, In: Water and Power in Past Societies (2018), Emily Holt, Proceedings of the IEMA Postdoctoral Visiting Scholar Conference on Theories and Methods in Archaeology, Vol. 6.

NOTE. You can download the chapter draft at Academia.edu.

Abstract (emphasis mine):

Along the western Norwegian coast, in the northwestern region of the Nordic Late Neolithic and Bronze Age (2350–500 BCE) there is cultural homogeneity but variable expressions of political hierarchy. Although new ideological institutions, technology (e.g., metallurgy and boat building), intensified agro‑pastoral farming, and maritime travel were introduced throughout the region as of 2350 BCE, concentrations of expressions of Bronze Age elites are intermittently found along the coast. Four regions—Lista, Jæren, Karmøy, and Sunnmøre—are examined in an exploration of the establishment and early role of maritime practices in this Nordic region. It is argued that the expressions of power and material wealth concentrated in these four regions is based on the control of bottlenecks, channels, portages, and harbors along important maritime routes of travel. As such, this article is a study of prehistoric travel, sources of power, and maritime landscapes in the Late Neolithic and Early Bronze Age of Norway.

Interesting excerpts:

(…)The [Corded Ware culture (CWC)] in Norway (or Battle Axe Culture, 2750–2400/2350 BCE) is primarily represented in Eastern Norway, with a patchy settlement pattern along the Oslo fjord’s coast through the inland valleys to Trøndelag in Central Norway (Hinsch 1956). The CWC represents an enigmatic period in Norwegian prehistory (Hinsch 1956; Østmo 1988:227–231; Prescott and Walderhaug 1995; Shetelig 1936); however the data at the moment suggests the following patterns:

  • Migration: The CWC was the result of a small‑scale immigration, but did not trigger substantial change.
  • Eastern and limited impact: The CWC was primarily located in small settlement patches in eastern Norway.
  • Terrestrial: In terms of maritime practices, the CWC does not represent a significant break from older traditions, though it seems to have a more pronounced terrestrial bearing. It is conceivable that pastures and hunting grounds were a more important political‑economic resource than waterways.

The mid‑third millennium in Norway, around 2400 BCE, represents a significant reorientation. Bell Beaker Culture (BBC) settlements in western Denmark and Norway archaeologically mark the instigation of the Nordic LN, though much of the historical process leading from the Bell Beaker to the Late Neolithic, 2500 to 2350 BCE, remains unclear (Prescott 2012; Prescott and Melheim 2009; Prieto‑Martinez 2008:116; Sarauw 2007:66; Vandkilde 2001, 2005). Still, the outcome is the establishment of the Nordic region of interaction in the Baltic, Northern Germany, Sweden, Denmark, and Norway. The distribution of artifact materials such as Bell Beakers and flint daggers attests to the far‑flung network of regular exchange and communication. This general region of interaction was reproduced through the Late Neolithic and Bronze Age.

nordic-late-neolithic
The Nordic region in the Late Neolithic and Bronze Age. Sites and regions discussed in the text are marked (ater Prescott and Glørstad 2015:fig. 1).

The transition from the preceding Neolithic period hunter‑gatherer societies was rapid and represents a dramatic termination of hunter‑gatherer traditions. It has been argued that the transformation is tied to initial migrations of people to the western coast of Norway from BBC areas, possibly from northern Jutland (Prescott 2011; Prescott and Walderhaug 1995:273). Bifacial tanged‑and‑barbed points, often referred to as “Bell Beaker points,” probably represent an early, short phase of the BBC‑transition around 2400 BCE. In Norway these points have a predominantly western and coastal distribution (Østmo 2012:64), underscoring the maritime nature of the initial BBC‑expansion.

late-neolithic-flint-daggers
Distribution routes for LN1 flint daggers type 1 suggesting communication routes and networks. (Redrawn after fig. 9, Apel 2001:17).

(…) In response to the question about what attracted people from Bell Beaker groups to western Norway, responses have hypothesized hunting products, political power, pastures, and metals. Particularly the latter has been emphasized by Lene Melheim (2012, 2015:37ff).

A recent study by Melheim and Prescott (2016) integrated maritime exploration with metal prospecting to explain initial excursions of BBC‑people along the western coast and into the fjords. Building on the archaeological concept of traveling metal prospectors as an element in the expansion of the Bell Beaker phenomenon, in combination with anthropological perspectives on prospecting, the article explores how prospecting for metal would have adjusted to the landscapes of western Scandinavia. Generally speaking, prospecting seldom leads to successful metal production, and it is difficult to study archaeologically. However, it will often create links between the prospectors’ society and indigenous groups, opening new territories, and have a significant transformative impact—on both the external and indigenous actors and societies.

While the text echoes the traditional idea that Corded Ware spread Indo-European languages, Prescott (since Prescott and Walderhaug 1995) is a supporter of the formation of a Nordic community and a Nordic (i.e. Pre-Germanic) language with the arrival of Bell Beakers.

An identification of the Corded Ware language as of a previous Proto-Indo-European stage is possible, as I have previously said (although my preference is Uralic-related languages).

This CWC language would thus still form the common substrate to both Germanic and Balto-Slavic, both being North-West Indo-European dialects, which spread with Bell Beakers over previous Corded Ware territory.

NOTE. This pre-LPIE nature could be in turn related to Kortlandt’s controversial proposal of an ealier PIE dative *-mus shared by both branches. However, that would paradoxically be against Kortlandt’s own assumption that the substrate was in fact of a non-Indo-European nature

See also:

Reproductive success among ancient Icelanders stratified by ancestry

iceland-pca

New paper (behind paywall), Ancient genomes from Iceland reveal the making of a human population, by Ebenesersdóttir et al. Science (2018) 360(6392):1028-1032.

Abstract and relevant excerpts (emphasis mine):

Opportunities to directly study the founding of a human population and its subsequent evolutionary history are rare. Using genome sequence data from 27 ancient Icelanders, we demonstrate that they are a combination of Norse, Gaelic, and admixed individuals. We further show that these ancient Icelanders are markedly more similar to their source populations in Scandinavia and the British-Irish Isles than to contemporary Icelanders, who have been shaped by 1100 years of extensive genetic drift. Finally, we report evidence of unequal contributions from the ancient founders to the contemporary Icelandic gene pool. These results provide detailed insights into the making of a human population that has proven extraordinarily useful for the discovery of genotype-phenotype associations.

icelanders
Shared drift of ancient and contemporary Icelanders. (A) Scatterplot of D-statistics reflecting Iceland-specific drift. To aid interpretation, we included values for ancient British-Irish Islanders and a subset of contemporary individuals (who were correspondingly removed from the reference populations).

We estimated the mean Norse ancestry of the settlement population (24 pre-Christians and one early Christian) as 0.566 [95% confidence interval (CI) 0.431–0.702], with a nonsignificant difference betweenmales (0.579) and females (0.521). Applying the same ADMIXTURE analysis to each of the 916 contemporary Icelanders, we obtained a mean Norse ancestry of 0.704 (95% CI 0.699–0.709). Although not statistically significant (t test p = 0.058), this difference is suggestive. A similar difference ofNorse ancestry was observed with a frequency-based weighted least-squares admixture estimator (16), 0.625 [Mean squared error (MSE) = 0.083] versus 0.74 (MSE = 0.0037). Finally, the D-statistic test D(YRI, X; Gaelic, Norse) also revealed a greater affinity between Norse and contemporary Icelanders (0.0004, 95% CI 0.00008–0.00072) than between Norse and ancient Icelanders (−0.0002, 95% CI −0.00056–0.00015). This observation raises the possibility that reproductive success among the earliest Icelanders was stratified by ancestry, as genetic drift alone is unlikely to systematically alter ancestry at thousands of independent loci (fig. S10). We note that many settlers of Gaelic ancestry came to Iceland as slaves, whose survival and freedom to reproduce is likely to have been constrained (17). Some shift in ancestry must also be due to later immigration from Denmark, which maintained colonial control over Iceland from 1380 to 1944 (for example, in 1930 there were 745 Danes out of a total population of 108,629 in Iceland) (18).

icelander-admixture
Shared drift of ancient and contemporary Icelanders. (B) Estimated Norse,
Gaelic, and Icelandic ancestry for ancient Icelanders using ADMIXTURE
in supervised mode.

Five pre-Christian Icelanders (VDP-A5, DAVA9, NNM-A1, SVK-A1 and TGS-A1) fall just outside the space occupied by contemporary Norse in Fig. 3A. That these individuals show a stronger signal of drift shared with contemporary Icelanders is also apparent in the results of ADMIXTURE, run in supervised mode with three contemporary reference populations (Norse, Gaelic, and Icelandic) (Fig. 3B). The correlation between the proportion of Icelandic ancestry from this analysis and PC1 in Fig. 2A is |r| = 0.913.(…)

(…) as the five ancient Icelanders fall well within the cluster of contemporary Scandinavians (Fig. 3C), we conclude that they, or close relatives, likely contributed more to the contemporary Icelandic gene pool than the other pre-Christians. We note that this observation is consistent with the inference that settlers of Norse ancestry had greater reproductive success than those of Gaelic ancestry.

icelanders-y-dna
Haplogroup data, from the paper. Image modified by me, with those close to Gaelic and British/Irish samples (see above Scatterplot of D-statistics and ADMIXTURE data) marked in fluorescent: yellow closer to Gaelic, green less close.

Ancient Icelanders show a clear relation with the typically Norse Y-DNA distribution: I1 / R1a-Z284 / R1b-U106.

  • Among R1a, the picture is uniformly of R1a-Z284 (at least five of the seven reported).
  • There are six samples of I1, with great variation in subclades.
  • Among R1b-L51 subclades (ten samples), there are U106 (at least one sample), L21 (three samples), and another P312 (L238); see above the relationship with those clustering closely with Gaelic samples, marked in fluorescent, which is compatible with Gaelic settlers (predominantly of R1b-L21 lineages) coming to Iceland as slaves.

Probably not much of a surprise, coming from Norse speakers, but they are another relevant reference for comparison with samples of East Germanic tribes, when they appear.

Also, the first reported Klinefelter (XXY) in ancient DNA (sample ID is YGS-B2).

Related:

The Indo-European demic diffusion model, and the “R1b – Indo-European” association

yamna_bell_beaker_cut

Beginning with the new year, I wanted to commit myself to some predictions, as I did last year, even though they constantly change with new data.

I recently read Proto-Indo-European homelands – ancient genetic clues at last?, by Edward Pegler, which is a good summary of the current state of the art in the Indo-European question for many geneticists – and thus a great example of how well Genetics can influence Indo-European studies, and how badly it can be used to interpret actual cultural events – although more time is necessary for some to realize it. Notice for example the distribution of ‘Yamnaya’ in 3000 BC, all the way to Latvia (based on the initial findings of Mathieson et al. 2017), and the map of 2000 BC with ‘Corded Ware’, both suggesting communities linked by admixture and unrelated to actual cultures.

Some people – especially those interested in keeping a simplistic picture of Europe, either divided into admixture groups or simplistic R1b-Vasconic / R1a-Indo-European / N1c-Uralic (or any combination thereof) – want (others) to believe that I am linking ‘Indo-Europeans’ with haplogroup R1b. That is simply not true. In fact, my model dismisses such simplistic identifications of the reconstructible proto-languages with any modern peoples, admixtures, or haplogroups.

vasconic-uralic
Simplistic Vasconic/R1b-Uralic/N1c distribution, and intruding Indo-European/R1a, according to Wiik.

The beauty of the model lies, therefore, precisely in that if you take any modern group speaking Indo-European languages, none can trace back their combination of language, admixture, and/or haplogroup to a common Indo-European-speaking people. All our ancestral lines have no doubt changed language families (and indeed cultures), they have admixed, and our European regions’ paternal lines have changed, so that any dreams of ‘purity’ or linguistic/cultural/regional continuity become absurd.

That conclusion, which should be obvious to all, has been denied for a long time in blogs and forums alike, and is behind the effort of many of those involved in amateur genetics.

Main linguistic aim

The main consequence of the model, as the title of the paper suggests, is that reconstructible Indo-European proto-languages expanded with people, i.e. with actual communities, which is what we can assert with the help of Genomics. From a personal (or ethnic, or political) point of view genomics is useless, but from an anthropological (and thus linguistic) point of view, genomics can be a very useful tool to decide between alternative models of language diffusion, which has given lots of headaches to those of us involved in Indo-European studies.

The demic diffusion theory for the three main stages of the proto-language expansion was originally, therefore, a dismissal of impossible-to-prove cultural diffusion models for the proto-language – e.g. the adoption of Late Proto-Indo-European by Corded Ware groups due to a patron-client relationship (as proposed by Anthony), or a long-lasting connection between cultures (as proposed by Kristiansen, and favoured by “constellation analogy” proponents like Clackson, who negated the existence of common proto-languages). It also means the acceptance of the easiest anthropological model for language change: migration and – consequently – replacement.

By the time of the famous 2015 papers, I had been dealing for some time with the idea that the shared features between Indo-Iranian and Balto-Slavic may have been due to a common substrate, and must have therefore had some reflection in genomic finds. The data on these papers, and the addition of a weak connection between Pre-Germanic and Balto-Slavic communities, together with their clearest genetic link – R1a-M417 subclades (especially European Z283) – made it still easier to propose a Corded Ware substrate, partially common to the three.

Allentoft Corded Ware
Allentoft et al. “Arrows indicate migrations — those from the Corded Ware reflect the evidence that people of this archaeological culture (or their relatives) were responsible for the spreading of Indo-European languages. All coloured boundaries are approximate.”

Before the famous 2015 papers (and even after them, if we followed their interpretation), we were left to wonder why the supposed vector of expansion of Indo-European languages, Corded Ware migrants – represented by R1a-Z645 subclades, and supposedly continued unchanged into modern populations in its ‘original’ ancestral territories, Balto-Slavic and Indo-Iranian – , were precisely the (phonetically) most divergent Indo-European languages – relative to the parent Late Indo-European proto-language.

My paper implied therefore the dismissal of an unlikely Indo-Slavonic group, as proposed by Kortlandt, and of a still less factible Germano-Slavonic, or Germano-Indo-Slavonic (?) group, as loosely implied by some in the past, and maybe supported in certain archaeological models (viz. Kristiansen or partially Anthony), and presently by some geneticists since their simplistic 2015 papers on “massive migrations from the steppe“, and amateur genetic fans with infinite pet theories, indeed.

A common Corded Ware substrate to Balto-Slavic and Indo-Iranian, and common also partially between Balto-Slavic and Germanic (as supported by Kortlandt, too, albeit with different linguistic connotations), would explain their common features. The Corded Ware culture (and Uralic, tentatively proposed by me as the group’s main language family) is a strong potential connection between them, further supported by phylogeography, too.

Other consequences

Interpretations in my paper help thus dismiss the simplistic Yamna -> Corded Ware -> Bell Beaker migration model implied with phylogeography in the 2000s, and revived again by geneticists and Kristiansen’s workgroup based on the famous 2015 papers, whereby – due to the “Yamnaya ancestral component” – the Yamna culture would have been composed of communities of R1a-M417 and R1b-M269 lineages which remained against all odds ‘related but separated’ for more than two thousand years, sharing a common unitary language (why? and how?), and which expanded from Yamna (mainly R1b-L23) into Corded Ware (mainly R1a-M417) and then into Bell Beaker (mainly R1b-L51), in imaginary migration waves whose traces Archaeology has not found, or Anthropology described, before.

While phylogeography (especially the distribution of ancient samples of certain R1b and R1a subclades) was the main genetic aspect I used in combination with Archaeology and Anthropology to challenge the reliability of the “Yamnaya ancestral component” in assessing migrations – and thus Kristiansen’s now-popular-again modified Kurgan model – , my main aim was to prove a recent expansion of Late Proto-Indo-European from the steppe, and a still more recent expansion of a common group of speakers of North-West Indo-European, the language ancestral to Italo-Celtic, Germanic, and probably Balto-Slavic (or ‘Temematic’, the NWIE substrate of Balto-Slavic, according to some linguists).

My arguments serve for this purpose, and modern distributions of haplogroups or admixture are fully irrelevant: I am ready to change my view at any time, regarding the role of any haplogroup, or ancestral component, archaeological data, or anthropological migration model, to the extent that it supports the soundest linguistic model.

proto-indo-european-stages
Stages of Proto-Indo-European evolution. IU: Indo-Uralic; PU: Proto-Uralic; PAn: Pre-Anatolian; PToch: Pre-Tocharian; Fin-Ugr: Finno-Ugric. The period between Balkan IE and Proto-Greek could be divided in two periods: an older one, called Proto-Greek (close to the time when NWIE was spoken), probably including Macedonian, and spoken somewhere in the Balkans; and a more recent one, called Mello-Greek, coinciding with the classically reconstructed Proto-Greek, already spoken in the Greek peninsula (West 2007). Similarly, the period between Northern Indo-European and North-West Indo-European could be divided, after the split of Pre-Tocharian, into a North-West Indo-European proper, during the expansion of Yamna to the west, and an Old European period, coinciding with the formation and expansion of the East Bell Beaker group.

Gimbutas’ old theory of sudden and recent expansion served well to support a real community of Proto-Indo-European speakers, as did later the Yamna -> Corded Ware -> Bell Beaker theory that circulated in the 2000s based on modern phylogeography, and as did later partially Anthony’s updated steppe theory (2007). On the other hand, Kristiansen’s long-lasting connections among north-west Pontic steppe cultures and Globular Amphorae and Trypillian cultures, did not fit well with a close community expanding rapidly – although recent genetic data on Trypillia and Globular Amphorae might be compelling him to improve his migration theory.

So, if data turns out to be not as I expect now, I will reflect that in future versions of the paper. I have no problem saying I am wrong. I have been wrong many times before, and something I am certain is that I am wrong now in many details, and I am going to be in the future.

If, for example, R1b-L23(xZ2105) is demonstrated to come from Hungary and not the steppe (as supported by Balanovsky) or R1a-M417 samples are proved to have expanded with West Yamna settlers (as recently proposed by Anthony, see below the Balto-Slavic question), I would support the same model from a linguistic point of view, but modified to reflect these facts. Or if a direct migration link is found in Archaeology from Yamna to Corded Ware, and from Corded Ware to Bell Beaker (as proposed in the 2015 papers), I will revise that too (again, see the image below). Or, if – as Lazaridis et al. (2017) paper on Minoans and Mycenaeans suggested – the Anatolian hypothesis (that is, one of the multiple ones proposed) turns out to be somehow right, I will support it.

calcolithic-expansion
My map of Late Proto-Indo-European expansion (A Grammar of Modern Indo-European, 2006), following Gimbutas and Mallory.

Haplogroups are the least important aspect of the whole model, they are just another data that has to be taken into account for a throrough explanation of migrations. It has become essential today because of the apparent lack of vision on the part of geneticists, who failed to use them to adjust their findings of admixture with findings of haplogroup expansions, favouring thus a marginal theory of long-lasting steppe expansion instead of the mainstream anthropological models.

Since many of these alternative scenarios seem less and less likely with each new paper, it is probably more efficient to talk about which developments are most likely to challenge my model.

Main points

My main predictions – based mostly on language guesstimates, archaeological cultures, and anthropological models of migration -, even with the scarce genomic data we had, have been proven right until know with new samples from Mathieson et al. (2017) and Olalde et al. (2017), among other papers of this past year. These were my original assumptions:

(1) A Middle Proto-Indo-European expansion defined by the appearance of steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1b-M269 and R1b-L23 lineages;

(2) A Late Proto-Indo-European expansion defined by steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1b-L23 subclades; and

(3) A North-West Indo-European expansion defined by steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1b-L51 subclades.

The expansion of Corded Ware peoples, associated with steppe ancestry + reduction in haplogroup diversity and expansion of (mainly) R1a-Z645 subclades, represents thus a different migration, which is compatible with the different nature of the Corded Ware culture, unrelated to Yamna and without migration waves from one to the other (although there were certainly contacts in neighbouring regions).

As you can see, neither of the 3+1 expansion models imply that no other haplogroup can be found in the culture or regions involved (others have in fact been found, and still the models remain valid): these migrations imply a reduction of haplogroup diversity, and the expansion of certain subclades as is common in population expansions throughout history. While we all accept this general idea, some people have difficulties accepting just those cases not compatible with their dreams of autochthonous continuity.

Nevertheless, there are still voids in genetic investigation.

Controversial aspects

In my humble opinion, these are potential conflict periods and the most likely areas of change for the future of the theory:

1. When and how did R1b-M269 lineages become “chiefs” in the steppe?

Based on scarce data from Khvalynsk, it seems that during the Neolithic there were many haplogroups in the North Pontic and North Caspian steppes. A reduction to R1b-M269 subclades must have happened either just before or (as I support) during (the migrations that caused) the Suvorovo-Novodanilovka expansion among Sredni Stog, probably coinciding also with the expansion (or one of the expansions) of CHG ancestry (and thus the appearance of ‘Steppe component’ in the steppe). My theory was based initially on Anthony’s account and TMRCA of haplogroups of modern populations (both ca. 4200-4000 BC), but recent samples of the Balkans (R1b-M269 and steppe ancestry) seem to trace the population expansion some centuries back.

If my assessment is correct, then modern populations of haplogroup R1b-M269* and R1b-L23* in the Balkans probably reflect that ancient expansion, and samples related to Proto-Anatolian cultures in the Balkans will most likely be of R1b-M269 subclades and R1b-L23*. After admixture in the Balkans, posterior migrations of Anatolian languages into Anatolia might be associated with a different admixture component and haplogroups, we don’t have enough data yet.

If the haplogroup reduction and expansion in Khvalynsk happened later than the Suvorovo-Novodanilovka expansion, then we might find the expansion of Pre- or Proto-Anatolian associated with many different haplogroups, such as R1b (xM269), R1a, I, J, or G2, and more or less associated with steppe ancestry in the Balkans.

Another reason for finding such variety of haplogroups in ancient samples from the Balkans would be that this Khvalynsk group of “chiefs” traversed – and mixed with – the Sredni Stog population. Nevertheless, if we suppose homogeneity in haplogroups in Khvalynsk during the expansion, a high proportion of different haplogroups explained by admixture with the local population of Sredni Stog would challenge the whole “chief domination” explanation by Anthony, and we would have to return to the “different culture” theory by Rassamakin and potentially an older migration from Khvalynsk. In any case, both researchers show clear links of the Suvorovo-Novodanilovka phenomenon to Khvalynsk, and a differentiation with the surrounding Sredni Stog culture.

A less likely model would support the identification of the whole Eneolithic Pontic-Caspian steppe as a loose Indo-Hittite-speaking community, which would be in my opinion too big a territory and too loose a cultural bond to justify such a long-lasting close linguistic connection. This will probably be the refuge of certain people looking desperately for R1a-IE connections. However, the nature of the western steppe will remain distinct from Late Proto-Indo-European, which must have developed in the Yamna culture, so autochthonous continuity is not on the table anymore, in any case…

suvorovo-novodanilovka-region
Coexistence of the Varna-Gumelniţa culture and the Suvorovo phase of the sceptre-bearer communities. 1 — Fălciu; 2 — Fundeni-Lungoţi; 3 — Novoselskaja; 4 — Suvorovo; 5 — Casimcea; 6 — Kjulevča; 7 — Reka Devnja; 8 — Drama; 9 — Gonova mogila; 10 — Reževo; 11 — geographically separate Decea variant of the sceptre bearer group (after Govedarica, Manzura 2011: Abb. 5, adapted).

2. How did R1a-M417 (and especially R1a-Z645) haplogroups came to dominate over the Corded Ware cultures?

If I am right (again, based on TMRCA of modern populations), then it is precisely at the time of the potential expansion of Proto-Corded Ware from the Dnieper-Dniester forest, forest-steppe, and steppe regions, ca 3300-3000. Furholt’s recent radiocarbon analysis and suggestions of a Lesser Poland origin of the third or A-horizon, on which disparate archaeologists such as Anthony or Klejn rely now, seem to suggest also that Corded Ware was a cultural complex rather than a compact culture reflecting a migration of peoples – similar thus to the Bell Beaker complex.

This cultural complex interpretation of Corded Ware contrasts with the quite homogeneous late samples we have, suggesting clear migration waves in northern Europe, at least at some point in time, so Genomics will be a great tool to ascertain when and from where approximately did Corded Ware peoples expand. Right now, it seems that Eneolithic Ukraine populations are the closest to its origin, so the traditional interpretation of its regional origin by Kristiansen or Anthony remains valid.

3. How was Indo-Iranian adopted by Corded Ware invaders?

This is rather an anthropological question. We need reasonable models of founder effect/cultural diffusion necessary for that to happen – similar to the ones necessary to explain the arrival of N1c subclades into north-east Europe, or the arrival of R1b subclades in Basque/Iberian-speaking regions in south-west Europe. My description of potential events in the eastern steppe – based partially on Anthony – is merely a short sketch. Genomic data is unlikely to offer more than it does today (replacement of haplogroups, and gradually of some steppe component, by late Corded Ware groups in the steppe), but let’s see what new samples can contribute.

As for what some Indians – and other people willing to confront them – are looking for, regarding R1a-M417 and/or Indo-European origins in India, I don’t see the point, we already know a) that the origin of the expansion is in the steppe and b) that Hindu nationalist biggots will not accept results from research that oppose their views. I don’t expect huge surprises there, just more fruitless discussions (fomented by those who live from trolling or conspiracies)…

4. Yamna settlers from Hungary

Anthony’s new theory – and the nature of Balto-Slavic – hinges on the presence of R1a-M417 subclades (associated with later Corded Ware samples) in Yamna settlers of Hungary, potentially originally from the North Pontic area, where the oldest sample has been found.

My ‘modified’ version of Anthony’s new model (the only I deem just remotely factible) includes the expansion of a Proto-Corded Ware from Lesser Poland, but (given the overwhelming R1b found in East Bell Beaker), with R1a-M417 being associated with the region. How to explain this language change with objective data? Well, we have Bell Beaker expanding to these areas at a later time, so we would need to find R1b-L23 settlers in Lesser Poland, and then a resurge of R1a-M417 haplogroup. If not, resorting yet again to cultural diffusion Yamna “patrons” to Corded Ware “clients” of Lesser Poland would bring us to square one, now with the ‘steppe ancestry’ controversy included…

Since some Eastern Europeans are (for no obvious reason whatsoever) putting their hopes on that IE-R1a-CWC association, let’s hope some samples of R1a-M417 in Yamna or Hungary give them a break, so that they can begin accepting something closer to mainstream anthropological models. We could then work from there a Yamna-> Bell Beaker / North-West Indo-European association truce, and from there keep accepting that no single haplogroup from Yamna settlers is linked with modern languages, cultures or ethnic groups.

yamna-region
localization of Central-European funerary monuments with elements of the Pit Grave culture (after Bátora 2006);

5. How and when was Balto-Slavic associated with haplogroup R1a?

If we accept the Southern or Graeco-Aryan nature of Balto-Slavic with influence from an absorbed North-West Indo-European dialect, “Temematic” (as Kortlandt does), then Indo-Slavonic adopted in the steppe from Potapovka by Sintashta and Poltavka populations divided ca. 2000 BC into Indo-Iranian (migrating to the east with Andronovo), and Balto-Slavic (migrating westward with the Srubna culture). History from there is not straightforward, and it should follow Srubna, Thraco-Cimmerian, or other late expansions from cultures of the steppe.

On the other hand, if it is a Northern dialect related closely to Germanic and Italo-Celtic (in a North-West Indo-European group), then its origin has to be found in the initial expansion of East Bell Beakers, and its development into either the Únětice culture (of Balkan and thus potentially “Southern IE” influence), or the Mierzanowice-Nitra culture (of Corded Ware and thus potentially Uralic influence), or maybe from both, given the intermediate substrate found in Germanic and Balto-Slavic.

It is my opinion that the association of Balto-Slavic with haplogroup R1a is quite early after the East Bell Beaker expansion, probably initially with the subclade typically associated with West Slavic, R1a-M458. I have not much data to support this (apart from the most common linguistic model), just modern haplogroup distribution maps and common TMRCA, and highly hypothetical archaeological-anthropological models. Genetics will hopefully bring more data.

Let’s see also what information on ancient haplogroups we can obtain from the Tollense valley (already showing a close cluster with modern West Slavic populations) and steppe regions.

6. How did Germanic, Celtic, and Italic expand?

Germanic is probably the most interesting one. Following the expansion of R1b-L51 subclades (especially R1b-U106) and steppe ancestry (a confounding factor, with the previous expansion of R1a-Z284 subclades) in Scandinavia is going to be fascinating. Anthropological models already point to a linguistic and archaeological expansion of Pre-Germanic with Bell Beaker peoples.

The expansion of Celtic seems to be associated with chiefdoms, untraceable today in terms of haplogroups, and it seems thus different from previous expansions. New studies might tell how that happened, if it was actually in successive ways, as proposed, or maybe we don’t have enough data yet to reach conclusions.

We don’t know either how Italic expanded into the Italian Peninsula, or whether Latin expanded with peoples from Italy, if at all, or it was mostly a cultural diffusion event, as it seems.

Regarding Etruscan, while I think it is a controversy initiated based on fantastic accounts, and ignited with few finds of Middle Eastern ancestry (that seem logical from the point of view of regional contacts), it will be important for Italian linguists and archaeologists, also to accept the most likely scenario.

As for Palaeo-Hispanic languages, while steppe ancestry is found quite reduced in R1b-L51 subclades (after so many different expansions and admixture events since the departure from the steppe), their distribution from the Chalcolithic onwards and the resurgence of native haplogroups may serve to ascertain which Pre-Roman tribes were associated with the oldest regions where these subclades dominated. For that aim, a closer look at the developments in Aquitania and other pre-Roman Vasconic- and Iberian-speaking regions may shed some light on how founder effects might develop to leave the native language intact (in a case similar to the adoption of Indo-Iranian by post-Corded Ware Sinthastha and Potapovka in the eastern Pontic-Caspian steppe).

NOTE: Although mostly unrelated, linguistic questions may also be somehow altered with a change of migration models. For example, our current Corded Ware Substrate Hypothesis – strongly contested by Kortlandt and others – implies that Uralic was potentially the language spoken by Eneolithic Ukraine / Proto-Corded Ware peoples, therefore early Uralic languages were spoken by Corded Ware peoples, as a substrate for Germanic and Balto-Slavic, and Balto-Slavic and Indo-Iranian. If an Indo-Hittite branch different from Late PIE is accepted for Eneolithic Ukraine (thus suggesting a millennia-long cultural-historical community in the steppe), then the model still stands (e.g. Ger. and BSl. *-mos/-mus, as stated by Kortlandt, would correspond to the oldest morphological IE layer). As you can read in the different versions of our model, the different possibilities for the common substrate are stated, and the most likely one selected. But the most likely a priori option sometimes turns out to be wrong…

NOTE 2: You can comment whatever you want here, but I opened a specific thread in our forum if you want serious comments on the model to stuck and be further discussed.

Featured images: from the book Interactions, changes and meanings. Essays in honour of Igor Manzura on the occasion of his 60th birthday. Țerna S., Govedarica B. (eds.). 2016. Kishinev: Stratum Plus.

See also: