R1a-Z280 and R1a-Z93 shared by ancient Finno-Ugric populations; N1c-Tat expanded with Micro-Altaic

Two important papers have appeared regarding the supposed link of Uralians with haplogroup N.

Avars of haplogroup N1c-Tat

Preprint Genetic insights into the social organisation of the Avar period elite in the 7th century AD Carpathian Basin, by Csáky et al. bioRxiv (2019).

Interesting excerpts (emphasis mine):

After 568 AD the Avars settled in the Carpathian Basin and founded the Avar Qaganate that was an important power in Central Europe until the 9th century. Part of the Avar society was probably of Asian origin, however the localisation of their homeland is hampered by the scarcity of historical and archaeological data.

Here, we study mitogenome and Y chromosomal STR variability of twenty-six individuals, a number of them representing a well-characterised elite group buried at the centre of the Carpathian Basin more than a century after the Avar conquest.

The Y-STR analyses of 17 males give evidence on a surprisingly homogeneous Y chromosomal composition. Y chromosomal STR profiles of 14 males could be assigned to haplogroup N-Tat (also N1a1-M46). N-Tat haplotype I was found in four males from Kunpeszér with identical alleles on at least nine loci. The full Y-STR haplotype I, reconstructed from AC17 with 17 detected STRs, is rare in our days. Only nine matches were found among haplotypes in YHRD database, such as samples from the Ural Region, Northern Europe (Estonia, Finland), and Western Alaska (Yupiks). We performed Median Joining (MJ) network analysis using N-Tat haplotypes with ten shared STR loci (Fig. 3, Table S9). All modern N-Tat samples included in the network had derived allele of L708 as well. Haplotype I (Cluster 1 in Fig. 3) is shared by eight populations on the MJ network among the 24 identical haplotypes. Cluster 1 represents the founding lineage, as it is described in Siberian populations, because this haplotype is shared by the most populations and it is more diverse than Cluster 2.

Nine males share N-Tat haplotype II (on a minimum of eight detected alleles), all of them buried in the Danube-Tisza Interfluve. We found 30 direct matches of this N-Tat haplotype II in the YHRD database, using the complete 17 STR Y-filer profile of AC1, AC12, AC14, AC15, AC19 samples. Most hits came from Mongolia (seven Buryats and one Khalkh) and from Russia (six Yakuts), but identical haplotypes also occur in China (five in Xinjiang and four in Inner Mongolia provinces). On the MJ network, this haplotype II is represented by Cluster 2 and is composed of 45 samples (including 32 Buryats) from six populations (Fig. 3).

y-str-haplogroup-n-mongolian-ugrians
Median Joining network of 162 N-Tat Y-STR haplotypes Allelic information of ten Y-STR loci were used for the network. Only those Avar samples were included, which had results for these ten Y-STR loci. The founder haplotype I (Cluster 1) is shared by eight populations including three Mongolian, three Székely, three northern Mansi, two southern Mansi, two Hungarian, eight Khanty, one Finn and two Avar (AC17, AC26) chromosomes. Haplotype II (Cluster 2) includes 45 haplotypes from six populations studied: 32 Buryats, two Mongolians, one Székely, one Uzbek, one Uzbek Madjar, two northern Mansi and six Avars (AC1, AC12, AC14, AC15, AC19 and KSZ 37). Haplotype III (indicated by a red arrow) is AC8. Information on the modern reference samples is seen in Table S9.

A third N-Tat lineage (type III) was represented only once in the Avar dataset (AC8), and has no direct modern parallels from the YHRD database. This haplotype on the MJ network (see red arrow in Fig. 3) seems to be a descendent from other haplotype cluster that is shared by three populations (two Buryat from Mongolia, three Khanty and one Northern Mansi samples). This haplotype cluster also differs one molecular step (locus DYS393) from haplotype II. We classified the Avar samples to downstream subgroup N-F4205 within the N-Tat haplogroup, based on the results of ours and Ilumäe et al.18 and constructed a second network (Fig. S4). The N-F4205 network results support the assumption that the N-Tat Avar samples belong to N-F4205 subgroup (see SI chapter 1d for more details).

Based on our calculation, the age of accumulated STR variance (TMRCA) within N-Tat lineage for all samples is 7.0 kya (95% CI: 4.9 – 9.2 kya), considering the core haplotype (Cluster 1) to be the founding lineage. Y haplogroup N-Tat was not detected by large scale Eurasian ancient DNA studies but it occurs in late Bronze Age Inner Mongolia and late medieval Yakuts, among them N-Tat has still the highest frequency.

Two males (AC4 and AC7) from the Transtisza group belong to two different haplotypes of Y-haplogroup Q1. Both Q1a-F1096 and Q1b-M346 haplotypes have neither direct nor one step neighbour matches in the worldwide YHRD database. A network of the Q1b-M346 haplotype shows that this male had a probable Altaian or South Siberian paternal genetic origin.

EDIT (5 APR 2019): The paper offers an interesting late sample before the arrival of Hungarian conquerors, although we don’t know which precise lineage the sample belongs to:

One sample in our dataset (HC9) comes from this population, and both his mtDNA (T1a1b) and Y chromosome (R1a) support Eastern European connections. (…) Furthermore, we excluded sample HC9 from population-genetic statistical analyses because it belongs to a later period (end of 7th – early 9th centuries)

Apparently, then, results are consistent with what was already known from studies of modern populations:

According to Ilumäe et al. study, the frequency peak of N-F4205 (N3a5-F4205) chromosomes is close to the Transbaikal region of Southern Siberia and Mongolia, and we conclude that most Avar N-Tat chromosomes probably originated from a common source population of people living in this area, completely in line with the results of Ilumäe et al.

haplogroup_n1
Geographic-Distribution Map of hg N3 from Ilumäe et al.

Finno-Ugrians share haplogroup R1a-Z280

Another paper, behind paywall, Genetic history of Bashkirian Mari and Southern Mansi ethnic groups in the Ural region, by Dudás et al. Molecular Genetics and Genomics (2019).

Interesting excerpts (emphasis mine):

Y‑chromosome diversity

The most frequent haplogroups of the Bashkirian Maris were N1b-P43 (42%), R1a-Z280 (16%), R1a-Z93 (16%), N1c-Tat (13%), and J2-M172 (7%). Furthermore, subgroup R1b-M343 accounted for 4% and I2a-P37 covered 2% of the lineages. None of the Mari N1c Y chromosomes belonged to the N1c subgroups investigated (L1034, VL29, Z1936).

In the case of the Southern Mansi males, the most frequent haplogroups were N1b-P43 (33%), N1c-L1034 (28%) and R1a-Z280 (19%). The frequencies of the remaining haplogroups were as follows: R1a-M458 (6%), I1-L22 (3%), I2a-P37 (3%), and R1b-P312 (3%). The haplotype and haplogroup diversities of the Bashkirian Mari group were 0.9929 and 0.7657, whereas these values for the Southern Mansi were 0.9984 and 0.7873, respectively. The results show that, in both populations, haplotypes are much more diverse than haplogroups.

bashkir-mari-southern-mansi
Haplogroup frequencies of the Bashkirian Mari and the Southern Mansi ethnic groups in Ural region

Genetic structure

(..) the studied Bashkirian Mari and Southern Mansi population groups formed a compact cluster along with two Khanty, Northern Mansi, Mari, and Estonian populations based on close Fst-genetic distances (< 0.05), with nonsignificant p values (p > 0.05) except for the Estonian population. All of these populations belong to the Finno-Ugric language family. Interestingly, the other Mansi population studied by Pimenoff et al. (2008) (pop # 38) was located a great distance from the Southern Mansi group (0.268). In addition, the Bashkir population (pop # 6) did not show a close genetic affinity to the Bashkirian Mari group (0.194), even though it is the host population. However, the Russian population from the Eastern European region of Russia (pop # 49) showed a genetic distance of 0.055 with the Southern Mansi group. All Hungarian speaking populations (pops 13, 22, 23, 24, 50, and 51) showed close genetic affinities to each other and to the neighbouring populations, but not to the two studied populations.

y-dna-hungarians-ugric-mansi
Multidimensional scaling (MDS) plot constructed on Fstgenetic distances of Y haplogroup frequencies of 63 populations compared. The haplogroup frequency data used for population comparison together with references are seen in Online Resource 2 (ESM_2). Pairwise Fst-genetic distances and p values between 63 populations were calculated as shown in Online Resource 3 (ESM_3) Fig. 4 Multidimensional scaling (MDS) plot constructed on Rstgenetic distances of 10 STR-based Y haplotype frequencies of 21 populations compared. Image modified to include labels of modern populations.

Phylogenetic analysis

Median-joining networks were constructed for:

N-P43 (earlier N1b):

(…) TMRCA estimates for this haplogroup were made for all P43 samples (n = 157) 8.7 kya (95% CI 6.7–10.8 kya), for the N-P43 Asian.

N1c-Tat:

(…) 75% of Buryats belonged to Haplotype 2, indicating that the Buryats studied by us is a young and isolated population (Bíró et al. 2015). Bashkirian Mari samples derive from Haplotype 2 via Haplotype 3 (see dark purple circles on the top of Fig. 6a). Haplotype 3 contained six males (2 Buryat, 1 Northern Mansi, and 3 Khanty samples from Pimenoff et al. 2008). The biggest Bashkirian Mari haplotype node (3 Mari samples) was positioned three mutational steps away from Haplotype 1 and the remaining Mari samples can be derived from this haplotype. Southern Mansi haplotypes were scattered within the network except for two, which formed a smaller haplotype node with two Northern Mansi and two Khanty samples from Pimenoff et al. (2008).

n1c-n-tat-uralic-ugric
Median-Joining Networks (MJ) of 153 N-Tat (a) and 26 N-L1034 (b) haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. For N-Tat network, we used data from Southern Mansi (n = 11), Bashkirian Mari (n = 6) samples with Hungarian (n = 12), Hungarian speaking Székely (n = 6), Northern Mansi (n = 14), Mongolian (n = 16), Buryat (n = 44), Finnish (n = 13), Uzbek Madjar (n = 2), Uzbek (n = 3), Khanty (n = 4) populations studied earlier by us (Fehér et al. 2015; Bíró et al. 2015) and Khanty (n = 18) and Mansi (n = 4) studied by Pimenoff et al. (2008)

R1a-Z280 haplotypes, shared by Maris, Mansis, and Hungarians, hence ancient Finno-Ugrians:

The founder R1a-Z280 haplotype was shared by four samples from four populations (1 Bashkirian Mari; 1 Southern Mansi; 1 Hungarian speaking Székely; and 1 Hungarian), as presented in Fig. 7 (Haplotype 1). Haplotype 2 included five males (3 Bashkirian Mari and 2 Hungarian), as it can be seen in Fig. 7. Haplotype 4 included two shared haplotypes (1 Bashkirian Mari and one Hungarian speaking Csángó). The remaining two Bashkirian Mari haplotypes differ from the founder haplotype (Haplotype 1) by two mutational steps via Hungarian or Hungarian and Bashkirian Mari shared haplotypes. Beside Haplotype 1, the remaining Southern Mansi haplotypes were shared with Hungarians (Haplotype 5 or turquoise blue and red-coloured circles above Haplotype 7) or with Hungarians and Hungarian speaking Székely group (Haplotypes 3, 5, and 6). Haplotype 7 included ten Hungarian speakers (Hungarian, Székely, and Csángó). One Hungarian and one Uzbek Khwarezm shared haplotype can be found in Fig. 7 as well (red and white-coloured circle). All the other haplotypes were scattered in the network. The age of accumulated STR variation within R1a-Z280 lineage for 93 samples is estimated to be 9.4 kya (95% CI 6.5–12.4 kya) considering Haplotype 1 (Fig. 7) to be the founder.

r1a-z280-ugrians
Median-Joining Networks (MJ) of 93 R1a-Z280 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used haplotype data from Bashkirian Mari (n = 7), Southern Mansi (n = 7), Hungarian (n = 52), Hungarian speaking Székely (n = 11), Hungarian speaking Csángó (n = 10), Uzbek Ferghana (n = 2), Uzbek Tashkent (n = 1), Uzbek Khwarezm (n = 1) and Northern Mansi (n = 2) populations

R1a-Z93 as isolated lineages among Permic and Ugric populations:

Figure 8 depicts an MJ network of R1a-Z93* samples using 106 haplotypes from the 14 populations (Fig. 8). All of the Bashkirian Mari samples (7 haplotypes) formed a very isolated branch and differed from the one Hungarian haplotype (Fig. 8, see Haplotype 1) by seven mutational steps as well from two Uzbek Tashkent samples (see Haplotype 3). Another Hungarian sample shared two haplotypes of Uzbek Khwarezm samples in Haplotype 4. This haplotype can be derived from Haplotype 3 (Uzbek Tashkent). Haplotype 2 included one Hungarian and one Khakassian male. The remaining three Hungarian haplotypes are outliers in the network and are not shared by any sample. The other population samples included in the network either form independent clusters such as Altaians, Khakassians, Khanties, and Uzbek Madjars or were scattered in the network. The age of accumulated STR variation (TMRCA) within R1a-Z93* lineage for 106 samples is estimated as 11.6 kya (95% CI 9.3–14.0 kya) considering an Armenian haplotype (Fig. 8, “A”) to be the founder and the median haplotype.

r1a-z93-ugrians
Median-Joining Networks (MJ) of 106 R1a-Z93 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used the next haplotype data: 7 Bashkirian Mari, 6 Khanty, 4 Uzbek Madjar, 5 Uzbek Ferghana, 9 Uzbek Tashkent, 7 Uzbek Khwarezm, 2 Mongolian, 2 Buryat, 6 Hungarian samples tested by us for this study or published earlier (Bíró et al. 2015) and populations (3 Armenian; 3 Afghan Tajik;
16 Altaian; 24 Khakassian; 12 Kyrgyz) from Underhill et al. (2015)

Comments

The results of modern populations for N (especially N1c) subclades show really wide clusters and ancient TMRCA, consistent with their known ancient and wide distribution in northern and eastern Eurasian groups, and thus with infiltration of different lineages with eastern nomads (and northern Arctic populations) coupled with later bottlenecks, as well as acculturation of groups.

EDIT (2 APR): Interesting is the specific subclade to which ancient Mongolic-speaking Avars belong (information from Yfull) N1c-F4205 (TMRCA ca. 500 BC), subclade of N1c-Y6058 (formed ca. 2800 BC, TMRCA ca. 2800 BC). This branch also gives the “European” branch N1c-CTS10760 (formed ca. 2800 BC, TMRCA ca. 2100 BC), and is subclade of a branch of N1c-L392 (formed ca. 4400 BC, TMRCA ca. 2800 BC). A northern expansion of N1c-L392 is probably represented by its branch N1c-Z1936 (formed ca. 2800, TMRCA ca. 2100 BC), the most likely candidate to appear in the Kola Peninsula in the Bronze Age as the Palaeo-Laplandic population (see here). Read more about potential routes of expansion of haplogroup N.

On the other hand, R1a-Z280 lineages form a tight cluster connecting Permic with Ugric groups, with R1a-Z93 showing early isolation (probably) between Cis-Urals and Trans-Urals regions. While both Corded Ware lineages in Finno-Ugrians are most likely related to the Abashevo expansion through Seima-Turbino and the Andronovo-like Horizon (and potentially later Eurasian expansions), a plausible hypothesis would be that Finno-Ugrians are related to an expansion of R1a-Z283 haplogroups (we already knew about the Finno-Permic connection), while the ancient connection between Permians and Hungarians with R1a-Z93 would correspond to this haplogroup’s potentially tighter link with an early Samoyedic split.

I don’t think that an explosive expansion of eastern Corded Ware groups of R1a-Z645 lineages will show a clear-cut division of haplogroups among Eastern Uralic groups, though, and culturally I doubt we will have such a clear image, either (similar to how the explosive expansion of Bell Beakers cannot be easily divided by regional/language group into R1b-L151 subclades before the known bottlenecks). Relevant in this regard are the known Z93 samples from the Árpád dynasty.

Nevertheless, this data may represent a slightly more recent wave of R1a-Z280 lineages linked to the expansion of Ugric into the Trans-Uralian region, after their split from Finno-Permic, still in close contact with Indo-Iranians in Poltavka and Sintashta-Potapovka, evident from the early and late Indo-Iranian borrowings, during a common period when Samoyedic had already separated.

Such a “Z283 over Z93” layer in the Trans-Urals (and Cis-Urals?) forest-steppes would be similar to the apparent replacement of Z284 by Z282 in the Eastern Baltic during the Bronze Age (possibly with the second or Estonian Battle Axe wave or, much more likely during later population movements). Such an early R1a-Z93 split could potentially be supported also by the separation into bottlenecks under “Northern” (R1a-Z283) Finno-Ugric-speaking Abashevo-related groups and “Southern” (R1a-Z93) acculturated Indo-Iranian-speaking Abashevo migrants developing Sintashta-Potapovka admixing with Poltavka R1b-Z2103 herders.

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

Conclusion

Let’s review some of the most common myths about Hungarians (and Finno-Ugrians in general) repeated ad nauseam, side by side with my assertions:

❌ N (especially N1c-Tat) in ancient and modern samples represent the True Uralic™ N1c peoples including Magyar tribes? Nope.

✅ Ancient N (especially N1c-Tat) lineages among Uralic populations expanded relatively recently, and differently in different regions (including eastern steppe nomads and northern arctic populations) not associated with a particular language or language group? Yep (read the series on Corded Ware = Uralic expansion).

❌ Modern Hungarian R1a-Z280 lineages represent the majority of the native population, poor Slavic ‘peasants’ from the Carpathian Basin, forcibly acculturated by a minority of bad bad Hungarian hordes? Nope.

✅ Modern Hungarian R1a-Z280 subclades represent Ugric lineages in common with ancient R1a-Z645 Finno-Ugric populations from north-eastern Europe and the Trans-Urals? Yep (see Avars and Ugrians).

❌ Modern Hungarian R1a-Z93 lineages represent acculturated Iranian/Turkic peoples from the steppes? Not likely.

✅ Modern Hungarian R1a-Z93 lineages represent a remnant of the expansion of Corded Ware to the east, potentially more clearly associated with Samoyedic? Much more likely.

finno-ugric-haplogroup-n
Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).

Sooo, the theory of a “diluted” Y-DNA in Modern Hungarians from originally fully N-dominated conquerors subjugating native R1a-Z280 Slavs from the Carpathian Basin is not backed up by genetic studies? The ethnic Iranian-Turkic R1a-Z93 federation in the steppes that ended up speaking Magyar is not real?? Who would’ve thunk.

Another true story whose rejection in genetics could not be predicted, like, not at all.

Totally unexpected, too, the drift of “R1a=IE” fans with the newest genetic findings towards a Molgen-like “Yamna/R1b = Vasconic-Caucasian”, “N1c = Uralic-Altaic”, and “R1a = the origin of the white world in Mother Russia”. So much for the supposed interest in “Steppe ancestry” and fancy statistics.

Related

15 thoughts on “R1a-Z280 and R1a-Z93 shared by ancient Finno-Ugric populations; N1c-Tat expanded with Micro-Altaic

  1. Magyars have miniscule % of N present in their y-dna, which in fact links them also to Siberian Uralic and also miniscule mtdna links to Siberian population. However, it does not rule out, that Magyar were already mixed R1a/N1a population when they were living on Ob river, where N and R were bordering each other. There is no reason to give up on probability, that females had main rule on passing language to their offsprings for some reason, or that uralic might have adopted and assimilated into their own tribe outsiders. Chechens had similar tepe(clans) of different ethnicities, which now are regarded as part of Chechens and they are indistinguishable from Chechens, because they speak the same language and wear the same national costume.
    It doesn’t mean, that initially male ancestors of those mixed children spoke Uralic, even if their mothers did so.
    Rest of Magyar y-dna(with some small variations) is indistinguishable from their neighbours – and just like their neighbours initially did not spoke Slavic, so something is very similar with Magyars, who clearly spoke some other languages before finally adapting Magyar. But it is most probable, that when Magyars arrived, they settled among Slavic speaking population.

    Uralic has ties with some other languages, that predates their contats with IE, so it is impossible, that any group of Uralic and any group belonging to y-dna N, who went out of their homeland located in China already spoke IE. Also Pit-Comb Ware of Uralic in Europe shows uninterrupted continuity from Pit-Comb Ware of Liao civilization. How that even makes Uralic as source for Corded-ware, if these were two distinct elements of different groups for a long time and who competed with each other up to the point, that R1a group slaughtered influx of migrants from north/east on Volga basin(at least that is what I’ve read in Russian about some charcoaled ruins, that were found on some river).

    If N1a-Tat expanded recently – even long after Corded-ware ended their expansion, how that makes Uralic as source for Corded-ware? And why? What is wrong with Uralic as Pit-Comb-ware? Even if we accept this controcversial claim, that Uralic was CWC, why there is no N1a-Tat in Germany and Scandinavia, where Corded-ware expanded? Btw, how are you going to explain Uralic in Baltic area, who never brought CWC from Finland area, but had their own distinct culture? No, dude – there were no Uralic CWC in Estonia and you are really confused, if you are trying to tie CWC with Uralic. CWC in Finland is Baltic or preBaltic – even most ancient hydronyms of Finland are Baltic. Saimaa lake in Finland is Baltic name and there are much bigger area of proposed hydronyms, which are only connectable to Baltic, but not Uralic.

    “not associated with a particular language or language group? Yep (read the series on”

    Actually, this is complete BS.

    Largest group of N1a nowadays are Russians and there are total ~10 million Russian males with N1a and their initial male ancestry is Uralic – even if they had Baltic ancestry, still – before that it was Uralic.
    Next largest group of 5-7 million total people is associated with Uralic speaking people, with exception of Selkups, who are mainly Q.
    Then the largest group is associated with Poles, Lithuanians, Latvians, Ukrainians, Gagauzi, who had ancient Uralic ancestry at some point and who are either coming from Novgorod assimilated Uralic, or Baltificied Uralic, who made into Lithuanian-Latvian and spread further.
    Then there are Swedes, who numbers in 1 million total, who have N1a male ancestry and that is either Saami ir Finn ancestry.
    Then there is another group of Tatars, Bashkirs or other Tukic people, who are very late arrivals in Siberia, who settled among Uralic, not to mention that only 500 years ago there were plentiful of Uralic tribes in Southern Siberia, who are now assimilated into Russians or others.
    Then there are Yakuts, which are brought up as an example by people who do not have any idea, that most of Yakut N1a ancestry is from assimilation of Yukagir, who were living before in Lena river basin. Some similar assimilations are happening to Dolgans and others who are slowly making into Yakut nation. And Yakuts are Nation, because they joined Russia as allies.

    Yukagirs spoke very similar language to Uralic, which actually is proposed as part of Yukagir-Uralic language group. So, no – it seems fit to father language hypothesis, just like it is in other cases, where if the language doesn’t fit to the group, then it also pokes out of current group.

    There are also some linguistic similarities that ties Uralic languages with Nivkhs and other paleo groups of Russian Far East. There are discussions about Samoyedic or Ugric languages as part of Uralic language group and that they make their own language group, but that does not mean, that they are not closer to Uralic than ANYTHING ELSE.

    So, no – most of N1a ancestry(except Eskimo, Aleutians, Chukchi and Japanese) is also linked with Yukaghir-Uralic. There are some other N groups and they actually do not have any links to distinct group, but that is not the case in N1a.

    It seems, that author has some major confusion about N1a in Baltic, which is not that complex problem for someone who is educated in this topic, but here are some hints for Baltic(which is my homeland, even though I have N1a and therefore not belonging to Baltic at all – because, why? why would someone insist on such lies?):
    1. There are no N1a in ancient samples up to 500BC(and in some places also 500AD) in Baltic. Virtually 100% of ancient Baltic is R1a and N1a is very recent(last 2000 years) addition in Baltic.
    2. Maps from Illumae are great, but they do not show, that Baltic had 2 distinct waves of N1a(they originated 2800BC-4000BC outside of Baltic) and could illuminate more about movements of N1a:
    1)CTS2929 which comes from east and is main composition of N in Lithuanian(90%).
    2)CTS10082(or other) which comes from north and is mostly present in Estonian.
    Unlike CTS10082, CTS2929 is probably mainly Uralic assimilated into Baltic with some BACKMIGRATION of Baltic colonists, along with their Baltic speaking relatives of Uralic male ancestry.

    It seems, that author has some major confusion how R1a has got into Uralic.
    Well, first of all – most ancient sample of R1a comes from Karelia ~8000BC and it predates anything with N1a. There were some other samples from Karelia, but it seems, that R1a came out as a winner.
    Then there was secondary expansion of R1a that went with Corded-ware and general direction for that expansion was towards north and east. Somehow I think, that it has to do with the rise of water levels in Black Sea, because Mediterranean connected with it in catastrophic event. Also, it is possible that at the same time there was Large Lake in the north of Siberia, which makes it impossible for people to live there at that time.
    Black Sea catastrophy is the main reason why R1a farmers went north. There was no reason to move north from Caspian Sea, which is not arable even today, because it is steppe located on ancient sea bed. So, ancient Mordvians(with Mescherians) and Mari(with Meryans) were safe from ancient nomads and your proposal is crap, because main assimilation of Uralic came from Baltic farmers – from West. Baltic left huge impact on Uralic – mainly agricultural and animal husbandry to mainly hunter-gatherer Uralic from whom Baltic gained a lot of words about hunting and fishing.

    Then there were more recent Baltic colonist movements into Volga basin out of their native Dniepr basin.
    Then there were some recent events, like Galindian colonists, who moved up to Moscow, who landed and assimilated local Baltic tribes(which is not shown in Illumae, but I assume, that Illumae refferred to something that predates event of Uralic assimilation by Baltic), then more recent Scandinavian-Prussian traders(1000AD), who established Novgorod, then Slavic movements, adoption of Church Slavonic along with orthodox etc. and not to mention that Russian south and Ukrainian east died out during Holodomor and replaced by Russians from central Russia and then we have some picture of final development of movements of people that are represented today.

    PS There are posted maps for R1a-M558… for some reason, which is 90% of R1a of Latvians. It is most Baltic R1a, because it overlaps areas where Baltic are living now and where Baltic lived in past(core area of Baltic was not near Baltic Sea). And even more – some of the R1a-M558 is present on area where nowadays Mari are located, even if there is assumption, that Baltic did not went so far. How the heck with that data did someone come to conclusion, that Balts = Uralic?
    I wonder why for example Mari =/= Baltic, because they have 50% R1a(unlike Baltic, who has it much less) and their R1a also seems to have high amounts of R1a-M558.

  2. Added information on the specific haplogroup of Mongolic-speaking Avars, N1c-F4205: it is well within the recent “European” N1c-L392 branch.

    An almost unreconstructible “Uralic-Altaic” family splitting ca. 2800 BC around Lake Baikal? Sure it is. Because haplogroup N.

  3. “On the other hand, R1a-Z280 lineages form a tight cluster connecting Permic with Ugric groups, with R1a-Z93 showing early isolation (probably) between Cis-Urals and Trans-Urals regions”

    I like this idea. Proto East-Uralic (Samoyedic and Ugric) splits off first and migrates to the east, admixing with the descendants of Yamnaya in the Volga-Ural region. This can account for the early appearance of R1a-Z93 in Potapovka c.2700 BCE (the oldest confirmed Z93) before the Abashevo time frame. Later, Abashevo expands with mostly R1a-Z283 lineages and spreading Finno-Permic dialects.

    I think I still favor a West Uralic (Finno-Permic) homeland in the mid-upper Volga and East Uralic (Ugric-Samoyedic, if that is possible) homeland in the Southern Urals during the time Proto Indo-Iranian was developing in the Volga-Ural and Trans-Ural steppes. This way, East Uralics will be the group directly admixing with Pre Indo-Iranians and the West Uralics will still be close to the action to account for all the IIr vocabulary in Finnic and Permic.

    1. [I think you mean Poltavka, although admittedly the attribution to this culture to an outlier from a shared cemetery with later samples may mean it was Potapovka]

      Yes, from what we know right now, it seems there will be two different bottlenecks, one of R1a-Z280 replacing only in part local I2a lineages with Abashevo, and one with R1a-Z93 in the steppes replacing R1b-Z2103. However, ‘archaic’ (I know, bad name) R1a-Z93 appears in Abashevo-related territory in the Trans-Urals (likely through Seima-Turbino) likely associated with Samoyedic, and probably also ‘archaic’ R1a-Z280 lineages will appear among Indo-Iranians in the EBA steppes, as we know already from Srubna.

      I think the main question for the Finno-Ugric homeland is whether or not these R1a-Z280 lineages proper of Abashevo really expanded with Netted Ware to the north-west into the Gulf of Finland. My bet is that they didn’t: we see textile ware everywhere since the Neolithic around the Baltic and in Eastern Europe and continued well into the BA, not related to a single culture or genetic community. I don’t doubt the genetic expansion of Textile Ware groups in NE Europe, but it seems the development around the Gulf of Finland was more local, and that must include the Finno-Samic community, while the so-called Netted Ware was different, and it may have been copied in some interaction regions. I haven’t seen much information published on these groups around Finland, maybe they are mostly written in Finnish. Indo-Iranian borrowings may be justified precisely through these contacts of Netted Ware with Estonian Textile Ware.

      In favor of your option, of Volga-Kama peoples expanding early to the north-west with Netted Ware, would be right now IMO the replacement of R1a-Z284 from the Baltic, haplogroups likely proper of Battle Axe, by mainly Z280 in subsequent migrations. On the other hand, the split of Z280 is so early that this replacement may have been due to migrations within the Baltic, i.e. related to the known shrinking and expansion of late Battle Axe-related BA groups, not necessarily to a broad population movement from the Volga. That’s one of the very interesting data to get from future papers on Abashevo and Fatyanovo-Balanovo.

      A real surprise could be the opposite, though: that R1a-Z280 expanded from Battle Axe / Fatyanovo-Balanovo to the south-east, replacing a R1a-Z93-dominated Abashevo. Sticking to the known models, Netted Ware seems to expand east to west, true, but the radiocarbon dates are too close, and the earliest textile ware in north-east Europe comes from the eastern Baltic, so let’s see if genetic movements upholds the proposal of Carpelan/Parpola, or if – as with the BBC – the theorized population movement must be reversed.

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