". Our resultssuggest that a new genetic component with strong Siberian affinity first arrivedin Europe around 4,000 years ago, as observed in our oldest analysedindividuals from northern Russia, and that the gene pool of modernnorth-eastern Europeans in general, and speakers of Uralic languages inparticular, is the result of multiple admixture events between Eastern andWestern sources since that first appearance. Additionally, we gain furtherinsights into the genetic history of the Saami in Finland, by showing that duringthe Iron Age, close genetic relatives of modern Saami lived in an area muchfurther south than their current geographic range."
"Most relevant to the populations analysed here is theadmixture cline between north-eastern Europe and the North SiberianNganasan, including mostly Uralic-speaking populations in our dataset (markedin light purple in Figure 2a).Ten of the eleven ancient individuals from this study fall on this Uralic cline, withthe exception of one individual from Levänluhta (JK2065), who is insteadprojected closer to modern Lithuanian, Norwegian and Icelandic populations.Specifically, two Levänluhta individuals and the two historical Saami fromRussia are projected very close to two previously published modern Saami(Saami.DG)23 and the new Saami genome generated in this study (as well as thepreviously published genome of the same individual, here labelled “Saami(WGA)”1), suggesting genetic continuity in the North from the Iron Age tomodern-day Saami populations. In contrast, the six ancient individuals fromBolshoy are projected much further towards East Asian populations, in anintermediate position along the Uralic cline and close to modern-day Mansi."
"The six ancient individuals from Bolshoy show substantially higherproportions of the Siberian component, which comprises about half of theirancestry (49.4-65.3 %), whereas the older Mesolithic individuals from Motala donot share this Siberian ancestry. The Siberian ancestry seen in EHG probably corresponds to a previously reported affinity towards Ancient North Eurasians(ANE)2,24, which also comprises part of the ancestry of Nganasans. Interestingly,results from uniparentally-inherited markers (mtDNA and Y chromosome) aswell as certain phenotypic SNPs also show Siberian signals in Bolshoy: mtDNAhaplogroups Z1, C4 and D4, common in modern Siberia18,25,26, in individualsBOO002, BOO004 and BOO006, respectively (confirming previous findings18), aswell as Y-chromosomal haplotype N1c1a1a (N-L392) in individuals BOO002 andBOO004. Haplogroup N1c, to which this haplotype belongs, is the major Ychromosomal lineage in modern North-East Europe and European Russia,especially in Uralic speakers, for example comprising as much as 54% of EasternFinnish male lineages today27. Notably, this is the earliest known occurrence ofY-haplogroup N1c in Fennoscandia. Additionally, within the Bolshoy populationwe observe the derived allele of rs3827760 in the EDAR gene, which is found innear-fixation in East Asian and Native American populations but is extremelyrare elsewhere28, and has been linked to phenotypes related to tooth shape29 andhair morphology30,31 (Supplementary Table 7). Scandinavian hunter-gatherersfrom Motala in Sweden have also been found to carry haplotypes associatedwith this allele4. Finally, we see high frequencies of haplotypes associated withdiets rich in high poly-unsaturated fatty acids, on the FADS genes4,32,33. The FADShaplotype observed here has previously been linked with Greenlanders32, and isfound in lower frequencies within Europe."
. To obtain a relative dateof this admixture, and as an independent line of evidence thereof, we usedadmixture linkage disequilibrium decay, as implemented in ALDER36. ALDERprovides a relative date estimate for a single-pulse admixture event ingenerations. When multiple admixture events have occurred, ALDER willprovide a date that is an average across the admixture processes36,37. Based onthe ALDER admixture date inferred for Bolshoy using Nganasan and EHG asadmixture sources, and assuming a generation time of 29 years38, we estimatethe time of introduction of Siberian ancestry in Bolshoy to 3,977 years beforepresent (yBP) (Figure 5b). Estimates obtained using Nganasan and Lithuaniansas source populations provided a similar estimate (Supplementary Figure 4 forLD decay plots for multiple populations using Lithuanians and Nganasan assources.). However, for all other populations with evidence for Siberianancestry, we find much younger admixture dates (Figure 5a), suggesting that theobserved genetic ancestry in north-eastern Europe is inconsistent with asingle-pulse admixture event. From the data shown in this study, the Bolshoyindividuals mark the earliest evidence of Siberian ancestry in north-easternEurope, while also predating the introduction of ancestry related to Neolithicfarmers into this area. As such, the admixture date estimate provided by ALDERfor Bolshoy is likely closer to the true time when Siberian ancestry was first in the area."
"The fact that the Siberian genetic component isconsistently shared among Uralic-speaking populations, with the exceptions ofHungarians and the non-Uralic speaking Russians, would make it tempting toequate this component with the spread of Uralic languages in the area. However,such a model may be overly simplistic. First, the presence of the Siberiancomponent on the Kola Peninsula at ca. 4000 yBP predates most linguisticestimates of the spread of Uralic languages to the area51. Second, as shown in ouranalyses, the admixture patterns found in historic and modern Uralic speakersare complex and in fact inconsistent with a single admixture event. Therefore,even if the Siberian genetic component partly spread alongside Uralic languages,it likely presented only an addition to populations carrying this component fromearlier.
"These results suggest thatthe geographic range of the Saami extended further south in the past, and hintsat a genetic shift at least in the western Finnish region during the Iron Age. Thefindings are in concordance with the noted linguistic shift from Saami languagesto early Finnish. Further ancient DNA from Finland is needed to conclude towhat extent these signals of migration and admixture are representative ofFinland as a whole."
So, they were probably there earlier.
