PL046 R-YP6228 PL048 I-PH833 PL049 I-A11537 PL052 R-Y48961 PL059 I-PH833 PL062 I-S15301 PL065 I-Y294193 PL066 R-FGC2555 PL067 R-S7759 PL070 I-CTS10028 PL071 I-BY316 PL076 I-S9318 PL082 I-Z2041 PL085 J-Z38241 PL086 I-FT29339See also... Early Slavs from Tribal Period Poland Wielbark Goths were overwhelmingly of Scandinavian origin High-resolution stuff
The only deviation from this pattern is observed for ancient samples from the Russian Volga-Oka region, where we measure higher genetic affinity between present-day Southern/Western Europeans and the SP population compared to the pre-SP population (Fig. S17). This agrees with the pattern observed in PCA and ADMIXTURE that, in contrast to the Northwestern Balkan, Eastern Germany, and Poland-Northwestern Ukraine, the arrival of Slavic-associated culture in Northwestern Russia was associated with a shift in PCA space to the West, a decrease of BAL [Baltic] ancestry, and the introduction of Western European ancestries such as CNE [Continental North European] and CWE [Continental Western European].Thus, it's highly plausible that the Tribal Period Slavs from Gródek were very similar, perhaps even practically identical, to the proto-Slavs who lived in the original Slavic homeland. Hopefully we won't have to wait too long to discover whether that's true or not. More Migration period and Slavic period samples from the border regions of Belarus, Poland and Ukraine are needed to sort that out. On the other hand, most of the post-1000 CE individuals from Gródek are shifted closer to present-day Balts. This is probably due to admixture from nearby Baltic-speaking populations. At the time, Baltic speakers still occupied much of northern and eastern Poland. I'm still going through the Gretzinger et al. paper and I'll probably have a lot more to say about it in the near future. However, unfortunately, I've already spotted a silly mistake in the supplementary info that will probably have some very annoying consequences for us on this blog. On page 109 the authors make the false claim that South Asian ancestry is present in a wide range of ancient Eastern European and Central Asian populations from the Bronze Age to the Scythian period.
Furthermore, Sycthian groups from Ukraine show varying fractions of South Asian ancestry (between 5% and 12%), a component present in many ancient individuals from Moldova (e.g. Moldova_IA, Moldova_LBA and Moldova_MBA), Ukraine (Ukraine_Alexandria_MBA and Ukraine_BA_Catacomb.SG), Western Russia (e.g. Russia_EarlySarmatian.SG, Russia_MLBA_Potapovka, or Russia_MLBA_Sintashta) and the Caucasus (Russia_Caucasus_LBA_Dolmen and Russia_North_Caucasus_MBA) but (nearly) absent in the SP genomes from Central and East-Central Europe (<5%) (Fig. S42b).All ancient and present-day South Asian populations carry what is commonly known as Ancestral South Indian (ASI) ancestry, while all of the above mentioned ancient groups lack it. Ergo, it's impossible for these ancients to have actual South Asian ancestry. What happened is that Gretzinger et al. created a genetic component in ADMIXTURE based on present-day South Asians. However, South Asians today have very complex ancestry from several different sources, including early pastoralists from the North-Pontic steppe in Eastern Europe and early farmers from Central Asia and what is now Iran. As a result, the groups that share significant amounts of alleles with South Asians via these sources also show so called South Asian ancestry in the Gretzinger et al. analysis. Unless this problem is corrected we're likely to see some nutjobs online using this paper to claim all sorts of nonsense about the origins of ancient Eastern Europeans and Central Asians, especially the Sintashta people and Scythians. See also... High-resolution stuff Leo Speidel & Pontus Skoglund
Popular methods of genetic analysis relying on allele frequencies such as PCA, ADMIXTURE and qpAdm are not suitable for distinguishing many populations that were important historical actors in the Migration Period Europe. For instance, differentiating Slavic, Germanic, and Celtic people is very difficult relying on these methods, but very helpful for archaeologists given a large proportion of graves with no inventory and frequent adoption of a different culture. To overcome these problems, we applied a method based on autosomal haplotypes. Imputation of missing genotypes and phasing was performed according to a protocol by Rubinacci et al. (2021), and IBD inference was done for ancient Eurasian individuals with data available at >600,000 1240K sites. IBD links for a subset of these individuals were represented as a graph, visualized with a force-directed layout algorithm, and clusters in this graph are inferred with the Leiden algorithm. One of the clusters in the IBD graph emerged that includes nearly all individuals in the dataset annotated archaeologically as “Slavic”. According to PCA a hypothesis for the origin of this population can be proposed: it was formed by admixture of a Baltic-related group with East Germanic people and Sarmatians or Scythians. The individuals belonging to the “Slavic” IBD sharing cluster form a chronological gradient on the PCA plot, with the earliest samples close to the Baltic LBA/EIA group. Later “Slavic” individuals are shifted to the right, closer to Central and Southern Europeans and probably reflecting further admixture of Slavs with local populations during the Migration Period.Apparently this abstract is causing a bit of confusion online because of the mention of possible Sarmatian or Scythian ancestry in Slavs. However, it's important to understand that the authors are referring to certain Slavic or even just Slavic-related individuals, usually from culturally heterogeneous frontier settlements deep in what is now Russia. So yes, it's possible that some of these individuals carry Sarmatian, Scythian or other exotic eastern ancestry. But even if this is true, then obviously we can't extend this inference to all ancient and modern-day Slavs. Indeed, below is a G25/Vahaduo Principal Component Analysis (PCA) that shows why modern-day Slavic speakers can't be linked genetically to Sarmatians or Scythians. To experience a more detailed version of the PCA paste the data here into the relevant field here.
Distance to: ITA_Sicily_Himera_480BCE_3:I10943 0.03393838 HUN_IA_La_Tene_o:I18226 0.03572886 DEU_MA_Krakauer_Berg:KRA001 0.03618075 RUS_Pskov_VA:VK159 0.03899963 SWE_Gotland_VA:VK463 0.03915018 Baltic_EST_IA:s19_V12_1 Distance to: ITA_Sicily_Himera_480BCE_3:I10949 0.03573636 HUN_IA_La_Tene_o3:I25524 0.03698768 HUN_IA_La_Tene_o:I18226 0.03732752 SWE_Skara_VA:VK397 0.03767022 Baltic_EST_IA:s19_V12_1 0.03772687 DEU_MA_Krakauer_Berg:KRA001On the other hand, I'm almost certain that the two Himera_480BCE_4 samples are Sarmatians. The good old G25 does it again!
Distance to: ITA_Sicily_Himera_480BCE_4:I10944 0.03100861 KAZ_Segizsay_Sarmatian:SGZ002 0.03548059 MDA_Sarmatian:I11925 0.03619219 RUS_Urals_Sarmatian:MJ56 0.03626538 RUS_Urals_Sarmatian:chy001 0.03904260 RUS_Urals_Sarmatian:MJ41 Distance to: ITA_Sicily_Himera_480BCE_4:I10947 0.02989458 RUS_Urals_Sarmatian:MJ43 0.03052790 RUS_Urals_Sarmatian:chy002 0.03170622 KAZ_Kangju:DA226 0.03288789 TUR_BlackSea_Samsun_Anc_C:I4529 0.03310149 KAZ_Aigyrly_Sarmatian:AIG003
- shared genetic drift with present-day Balto-Slavic speakers to the exclusion of most other Europeans - and yet, an unusually low level of Yamnaya-related steppe ancestry - so much so, in fact, that they're often outside the range of modern European genetic variation.As far as I can tell, currently the best examples of this unusual population are HUN_Mako_EBA_o:I1502 (Mathieson et al. Nature 2015) and HUN_EIA_Prescythian_Mezocsat_o1:I18241 (Patterson et al. Nature 2021). Both are from the Carpathian Basin in what is now Hungary. I ran a series of qpAdm mixture models to try and learn more about their origins. The most robust outcomes, out of about 50 different attempts, are these:
right pops: CMR_Shum_Laka_8000BP MAR_Taforalt IRN_Ganj_Dareh_N Levant_PPNB TUR_Barcin_N Iberia_Southeast_Meso UKR_Meso England_Meso RUS_Karelia_HG RUS_West_Siberia_HG MNG_North_N TWN_Hanben BRA_LapaDoSanto_9600BP HUN_Mako_EBA_o Baltic_LTU_Narva 0.149 ∓0.028 POL_Globular_Amphora 0.613 ∓0.028 Yamnaya_RUS_Samara 0.238 ∓0.029 chisq 10.836 tail prob 0.370463 Full output HUN_EIA_Prescythian_Mezocsat_o1 Baltic_LTU_Narva 0.186 ∓0.028 POL_Globular_Amphora 0.592 ∓0.027 Yamnaya_RUS_Samara 0.222 ∓0.029 chisq 12.492 tail prob 0.253499 Full outputCombining the two genomes produces a very similar result:
HUN_EBA-EIA_o Baltic_LTU_Narva 0.160 ∓0.023 POL_Globular_Amphora 0.612 ∓0.023 Yamnaya_RUS_Samara 0.227 ∓0.023 chisq 14.653 tail prob 0.14524 Full outputImportantly, when I move RUS_Karelia_HG from the right pops to the left pops, to test whether HUN_EBA-EIA_o really has steppe ancestry, as opposed to closely related hunter-gatherer ancestry, I still get a very similar outcome:
HUN_EBA-EIA_o Baltic_LTU_Narva 0.158 ∓0.027 POL_Globular_Amphora 0.605 ∓0.033 RUS_Karelia_HG 0.014 ∓0.038 Yamnaya_RUS_Samara 0.223 ∓0.053 chisq 10.461 tail prob 0.234171 Full outputSo these largely Globular Amphora-related individuals do harbor as much as a quarter of steppe ancestry, which is to be expected considering the massive genetic turn-over that most of Europe experienced just before their time as a result of population expansions from the Pontic-Caspian steppe. Nevertheless, this is ~20% less steppe ancestry than in the present-day populations of the region, and it clearly shows in any decent Principal Component Analysis (PCA) of West Eurasia. For instance:
In addition to autosomal genetic changes through time, we observe a sharp reduction in Y-chromosomal diversity going from five different lineages in early CW to a dominant (single) lineage in late CW (Fig. 4A). We used forward simulations to explore the demographic scenarios that could account for the observed reduction in Y-chromosomal diversity. Performing 1 million simulations of a population with a starting frequency of R1a-M417(xZ645) centered around the observed starting frequency in Bohemia_CW_Early (3 of 11, 0.27), we assessed the plausibility of this lineage reaching the observed frequency in Bohemia_CW_Late (10 of 11, 0.91) in the time frame of 500 years under a model of a closed population and random mating (Materials and Methods). We reject the “neutral” hypothesis, i.e., that this change in frequency occurred by chance, given a wide range of plausible population sizes. Instead, our results suggest that R1a-M417(xZ645) was subject to a nonrandom increase in frequency, resulting in these males having 15.79% (4.12 to 44.42%) more surviving offspring per generation relative to males of other Y-haplogroups. We also find that this change in Y chromosome frequency is extreme compared to the changes in allele frequencies at fully covered autosomal 1240k sites within the same males, suggesting a process that disproportionately affected Y-chromosomal compared to autosomal genetic diversity, ruling out a population bottleneck as the likely cause. Our results suggest that the Y-lineage diversity in early CW males was supplanted by a nonrandom process [selection, social structure, or influx of nonlocal R1a-M417(xZ645) lineages] that drove the collapse in Y-chromosomal diversity. A simultaneous decline of Y-chromosomal diversity dating to the Neolithic has been observed across most extant Y-haplogroups (64), possibly due to increased conflict between male-mediated patrilines (65). We view that changes in social structure (e.g., an isolated mating network with strictly exclusive social norms) could be an alternative cause but would be difficult to distinguish in the underlying model parameters.Right, so even though the CWC was clearly a community of closely related groups, there must have been some competition between its different clans. And since these clans were highly patriarchal and patrilineal, this competition probably led to different paternal lineages dominating different parts of the CWC horizon, with M417 becoming especially common in the east and L51 in the west. Of course, the expansions of post-Corded Ware groups, such as the M417-rich Slavs in Eastern Europe and L51-rich Celts in Western Europe, were also instrumental in creating Europe's R1a/R1b dichotomy, but obviously these groups were in large part the heirs of the CWC. By the way, most of the samples from Papac et al. are already in the Global25 datasheets linked here. Look for the labels listed here. Below is a plot made from the Global25 data courtesy of regular commentator Matt.
David, when are you going to explain the genetic discrepancy between Northeastern and Northwestern Europeans? You know, the one that people believe is due to Baltic Hunter-Gatherer admixture, whereas you believe it is due to genetic drift? You ought to make a post about this issue at some point, because a lot of people are wondering what's causing the differences.Well, Greg, this issue has been discussed to the proverbial death here and elsewhere. In fact, there were two posts and rather lengthy comment threads on the same topic at this blog just a few months ago. See here and here. Nevertheless, it seems that a fair number of people are still befuddled, so I'm going to try to explain this one last time, as briefly as a I can using just a handful of f4-stats. Admittedly, Northeast Europeans generally do pack higher levels of indigenous European hunter-gatherer ancestry than Northwest Europeans. This is especially true of Balts, who show more of this type of ancestry than even Scandinavians in practically every type of analysis. The f4-stats below back this up unambiguously. Note the significantly positive (>3) Z scores, which suggest that Latvians and Lithuanians harbor more Baltic hunter-gatherer-related ancestry than Norwegians and Swedes.
Chimp Baltic_HG Norwegian Latvian 0.001301 7.114 Chimp Baltic_HG Swedish Latvian 0.001017 4.205 Chimp Baltic_HG Norwegian Lithuanian 0.001023 7.341 Chimp Baltic_HG Swedish Lithuanian 0.000763 3.408Greg, I know what you're thinking: the naysayers are right! But wait, because there's a twist to this tale. Check out these f4-stats:
Chimp Baltic_HG Norwegian Belarusian 0.000265 1.934 Chimp Baltic_HG Swedish Belarusian 0.000152 0.7 Chimp Baltic_HG Norwegian Polish 6.4E-05 0.519 Chimp Baltic_HG Swedish Polish -0.000235 -1.074Please note, Greg, that none of the Z scores reach significance, which means that these Northwest Europeans and Slavs are symmetrically related to Baltic_HG. They're also symmetrically related to other relevant ancient groups such as the Yamnaya steppe herders. This, of course, suggests that they harbor very similar levels of basically the same ancient genetic components.
Chimp Karelia_HG Norwegian Belarusian 0.000136 0.844 Chimp Karelia_HG Swedish Belarusian 7.9E-05 0.32 Chimp Karelia_HG Norwegian Polish -4.7E-05 -0.304 Chimp Karelia_HG Swedish Polish -0.000134 -0.54 Chimp Yamnaya_Samara Norwegian Belarusian -0.000134 -1.085 Chimp Yamnaya_Samara Swedish Belarusian -6.6E-05 -0.34 Chimp Yamnaya_Samara Norwegian Polish -0.000225 -1.995 Chimp Yamnaya_Samara Swedish Polish -0.000311 -1.574 Chimp Barcin_N Norwegian Belarusian -0.000335 -2.809 Chimp Barcin_N Swedish Belarusian -0.000284 -1.491 Chimp Barcin_N Norwegian Polish -0.000222 -2.057 Chimp Barcin_N Swedish Polish -0.000318 -1.662 Chimp Baikal_N Norwegian Belarusian 0.000186 1.3 Chimp Baikal_N Swedish Belarusian -7E-05 -0.33 Chimp Baikal_N Norwegian Polish -4.6E-05 -0.351 Chimp Baikal_N Swedish Polish -0.000477 -2.277Interestingly, pairing up Ukrainians with English samples from Cornwall and Kent produces similar outcomes. But that's because most ancient ancestry proportions in Europe show a closer correlation with latitude than longitude.
Chimp Baltic_HG English_Cornwall Ukrainian 0.000282 2.242 Chimp Baltic_HG English_Kent Ukrainian 0.000225 1.748 Chimp Karelia_HG English_Cornwall Ukrainian 0.000323 2.175 Chimp Karelia_HG English_Kent Ukrainian 0.000239 1.634 Chimp Yamnaya_Samara English_Cornwall Ukrainian -6.6E-05 -0.569 Chimp Yamnaya_Samara English_Kent Ukrainian -0.000112 -0.977 Chimp Barcin_N English_Cornwall Ukrainian -0.000519 -4.641 Chimp Barcin_N English_Kent Ukrainian -0.000598 -5.232 Chimp Baikal_N English_Cornwall Ukrainian 0.000385 2.874 Chimp Baikal_N English_Kent Ukrainian 0.00036 2.836Now, Greg, if at least in terms of genetic ancestry, Latvians, Lithuanians, Belarusians, Poles and Ukrainians all qualify as Northeast Europeans, then what makes them different, as a group, from Northwest Europeans? Do you believe that the key factor is admixture from Baltic hunter-gatherers? Or is it genetic drift? Of course, considering all of the f4-stats above, logic dictates that it must be relatively recent genetic drift. Keep in mind, however, that this only applies to Balto-Slavic speaking Northeast Europeans without significant Uralian ancestry. Overall, Uralic speakers have a more complex population history, and indeed genetic differences between them and Northwest Europeans are in large part due to somewhat different ancestry proportions and also Siberian admixture. See also... So who's the most (indigenous) European of us all?
In the traditional narrative, the Fatyanovo people – like the CWC populations in general – are regarded as Indo-European, representing the pre-Balto-Slavic (-Germanic) stage (Carpelan & Parpola 2001, 88; Anthony 2007, 380; also Gimbutas 1956, 163; Tretyakov 1966, 109) in the spread of Indo-European languages.That's correct, but considering the latest ancient DNA research on the Fatyanovo people, the traditional narrative is probably wrong. Fatyanovo males were rich in Y-haplogroup R1a-Z93, which is found at very low frequencies in Balto-Slavic populations (see here). It's actually much more common nowadays in Central and South Asia, where it often reaches frequencies of over 50% in Indo-Iranian speaking groups. Balts and Slavs are rich in R1a-Z282, which is a sister clade of R1a-Z93 that has been found in Corded Ware and Corded Ware-related samples from west of Fatyanovo sites. That is, in present-day Poland and the Baltic states. Therefore, the origins of the Balto-Slavs should be sought somewhere west of the Fatyanovo culture, probably in the Corded Ware derived populations from what is now the border zone between Poland, Belarus and Ukraine. Indeed, in my view the Fatyanovo people are more likely to have spoken Proto-Indo-Iranian rather than anything ancestral to Baltic or Slavic (see here).
- this may have been a battle between two Central European armies, one rich in Y-haplogroup R1b and the other rich in Y-haplogroup I2a, as well as their allies or hired help, including warriors from Eastern Europe belonging to Y-haplogroup R1a - or perhaps it was an invasion from the east by warriors rich in Y-haplogroup R1a, and it was a success, with the local armies, rich in Y-haplogroups R1b and I2a, losing the battle and suffering most of the casualties.I'm sure that one day someone will attempt to undertake a decent multidisciplinary study of this epic battle, and we'll at least have a rough idea about what happened. Or not. Citation... Burger et al., Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years, Current Biology, Available online 3 September 2020, https://doi.org/10.1016/j.cub.2020.08.033 See also... Genetic and linguistic structure across space and time in Northern Europe
Lactase persistence (LP), the continued expression of lactase into adulthood, is the most strongly selected single gene trait over the last 10,000 years in multiple human populations. It has been posited that the primary allele causing LP among Eurasians, rs4988235-A [1], only rose to appreciable frequencies during the Bronze and Iron Ages [2, 3], long after humans started consuming milk from domesticated animals. This rapid rise has been attributed to an influx of people from the Pontic-Caspian steppe that began around 5,000 years ago [4, 5]. We investigate the spatiotemporal spread of LP through an analysis of 14 warriors from the Tollense Bronze Age battlefield in northern Germany (∼3,200 before present, BP), the oldest large-scale conflict site north of the Alps. Genetic data indicate that these individuals represent a single unstructured Central/Northern European population. We complemented these data with genotypes of 18 individuals from the Bronze Age site Mokrin in Serbia (∼4,100 to ∼3,700 BP) and 37 individuals from Eastern Europe and the Pontic-Caspian Steppe region, predating both Bronze Age sites (∼5,980 to ∼3,980 BP). We infer low LP in all three regions, i.e., in northern Germany and South-eastern and Eastern Europe, suggesting that the surge of rs4988235 in Central and Northern Europe was unlikely caused by Steppe expansions. We estimate a selection coefficient of 0.06 and conclude that the selection was ongoing in various parts of Europe over the last 3,000 years.Burger et al., Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years, Current Biology, Available online 3 September 2020, https://doi.org/10.1016/j.cub.2020.08.033 See also... Warriors from at least two different populations fought in the Tollense Valley battle
Transition from the Stone to the Bronze Age in Central and Western Europe was a period of major population movements originating from the Ponto-Caspian Steppe. Here, we report new genome-wide sequence data from 28 individuals from the territory north of this source area - from the under-studied Western part of present-day Russia, including Stone Age hunter-gatherers (10,800-4,250 cal BC) and Bronze Age farmers from the Corded Ware complex called Fatyanovo Culture (2,900-2,050 cal BC). We show that Eastern hunter-gatherer ancestry was present in Northwestern Russia already from around 10,000 BC. Furthermore, we see a clear change in ancestry with the arrival of farming - the Fatyanovo Culture individuals were genetically similar to other Corded Ware cultures, carrying a mixture of Steppe and European early farmer ancestry and thus likely originating from a fast migration towards the northeast from somewhere in the vicinity of modern-day Ukraine, which is the closest area where these ancestries coexisted from around 3,000 BC. ... Interestingly, in all individuals for which the chrY hg could be determined with more depth (n=6), it was R1a2-Z93 (Table 1, Supplementary Data 2), a lineage now spread in Central and South Asia, rather than the R1a1-Z283 lineage that is common in Europe [38,39].
