The arrival of N1a1-M46 (previously called N1c1) lineages into Northern Europe has been dated after 5000 BC[Ilumae et al. 2016], coinciding with the TMRCA of N1a1a1a-L708. However, the more recent formation (ca. 3800 BC) and TMRCA (ca. 2300 BC) of common European lineage N1a1a1a1a-L392 point to a late and stepped spread of these hunter-gatherer groups into the Forest Zone, that cannot be associated with the expansion of Pit-Comb Ware cultures.
Cultural assimilation remains therefore the best explanation for the shared Uralic languages of modern communities with a majority of R1a1a1-M417 and N1a1-M46 lineages. Eastern groups with N1a1a1a1a-L392 lineages may have brought with them the Altaic traits found in Uralic languages[Kortlandt 2010].
An aDNA sample of haplogroup N1a-F1206 is found in the Forest Zone dated ca. 2500 BC at Serteya[Chekunova et al. 2014]. Nevertheless, it is tempting to place the mass migration of Siberian hunter-gatherer communities around the Urals later, with the expansion of the poorly understood Seima-Turbino phenomenon (which began ca. 2000 BC in East Asia), since it connected cultures from Mongolia to Finland.
Three samples of haplogroup NO (xO) found in the Middle Bronze Age Okunev culture, and two samples later in the Chermuchek culture area[Hollard et al. 2014] may give support to this assumption.
Modern Estonians have R1a-M417 lineages in more than a third of their population[Laitinen et al. 2002], similar to the proportion of haplogroup N1a1-M46[Rosser et al. 2000]. However, Finns show almost two thirds of haplogroup N1a1-M46 in the modern population, and only about 10% of R1a-M417.
Estonians are close genetic relatives to Finns, as well as to Baltic peoples and Russians from the Tver region[Nelis et al. 2010]. Investigation of mtDNA in modern Finns has shown that there was probably a population decline ca. 1500 BC, and later an Iron Age bottleneck with a population peak ca. 500 AD[Översti et al. 2017].
There was probably, then, a long-term, gradual replacement of previously prevalent Y-DNA R1a-M417 subclades in the region, as supported by the increased ‘steppe’ ancestry in genome-wide ancestry of modern Finns. A sudden, strong population (and cultural) change associated with the arrival of N1a1-M46 lineages – like the ones seen with R1a-M417 (Corded Ware) and R1b-M269 (Yamna) in eastern Europe, has to be rejected. A founder effect of N1a1-M46 lineages is therefore the most likely explanation for their adoption of Finnic languages, a situation that is also supported by the genetic diversity of the Saami population[Tambets et al. 2004].
- [Anthony 2007] ^ Anthony, D. 2007. The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World. Princeton and Oxford: Princeton University Press.
- [Chekunova et al. 2014] ^ Chekunova, Е.М., N.V. Yartseva, М.К. Chekunov, and А.N. Мazurkevich. 2014. The First Results of the Genotyping of the Aboriginals and Human Bone Remains of the Archeological Memorials of the Upper Podvin’e. // Archeology of the lake settlements of IV—II Thousands BC: The chronology of cultures and natural environment and climatic rhythms. Paper read at Proceedings of the International Conference, Devoted to the 50-year Research of the Pile Settlements on the North-West of Russia., 13-15 November, at St. Petersburg.
- [Fokkens and Harding 2013] ^ Fokkens, Harry, and Anthony Harding. 2013. The Oxford Handbook of European Bronze Age. Oxford: Oxford University Press.
- [Hanks, Epimakhov, and Renfrew 2015] ^ Hanks, B. K., A. V. Epimakhov, and A. C. Renfrew. 2015. Towards a refined chronology for the Bronze Age of the southern Urals, Russia. Antiquity 81 (312):353-367.
- [Hollard et al. 2014] ^ Hollard, C., C. Keyser, P. H. Giscard, T. Tsagaan, N. Bayarkhuu, J. Bemmann, E. Crubezy, and B. Ludes. 2014. Strong genetic admixture in the Altai at the Middle Bronze Age revealed by uniparental and ancestry informative markers. Forensic Sci Int Genet 12:199-207.
- [Kortlandt 2010] ^ Kortlandt, Frederik 2010. Indo-Uralic and Altaic revisited. In Transeurasian verbal morphology in a comparative perspective: genealogy, contact, chance. Wiesbaden: Harrassowitz.
- [Ilumae et al. 2016] ^ Ilumae, A. M., M. Reidla, M. Chukhryaeva, M. Jarve, H. Post, M. Karmin, L. Saag, A. Agdzhoyan, A. Kushniarevich, S. Litvinov, N. Ekomasova, K. Tambets, E. Metspalu, R. Khusainova, B. Yunusbayev, E. K. Khusnutdinova, L. P. Osipova, S. Fedorova, O. Utevska, S. Koshel, E. Balanovska, D. M. Behar, O. Balanovsky, T. Kivisild, P. A. Underhill, R. Villems, and S. Rootsi. 2016. Human Y Chromosome Haplogroup N: A Non-trivial Time-Resolved Phylogeography that Cuts across Language Families. Am J Hum Genet 99 (1):163-73.
- [Jaeger 2012] ^ Jaeger, Mateusz. 2012. Enclosed Space - Open Society, Contact and Exchange in the Context of Bronze Age Defensive Settlements in Central Europe. In SAO/SPES, edited by M. Jaeger, J. Czebreszuk and K. P. Fischl. Poznań/Bonn: Bogucki Wydawnictwo Naukowe / Dr. Rudolf Habelt GmbH.
- [Krause 2013] ^ Krause, Rüdiger. 2013. Die bronzezeitliche Sintašta-Kultur im Trans-Ural – Impulsgeber in der Eurasischen Steppe. In Landschaft, Besiedlung und Siedlung. Archäologische Studien im nordeuropäischen Kontext, edited by I. Heske, H.-J. Nüsse and J. Schneeweiss. Neumünster/Hamburg: Wachholtz.
- [Kristiansen and Larsson 2005] ^ Kristiansen, Kristian, and Thomas B. Larsson. 2005. The Rise of Bronze Age Society: Travels, Transmissions and Transformations. Cambridge: Cambridge University Press.
- [Laitinen et al. 2002] ^ Laitinen, V., P. Lahermo, P. Sistonen, and M. L. Savontaus. 2002. Y-Chromosomal Diversity Suggests that Baltic Males Share Common Finno-Ugric-Speaking Forefathers. Human Heredity 53 (2):68-78.
- [Meller et al. 2015] ^ Meller, Harald, Helge Wolfgang Arz, Reinhard Jung, and Roberto Risch. 2015. 2200 BC - A climatic breakdown as a cause for the collapse of the old world? 7th Archaeological Conference of Central Germany October 23-26, 2014 in Halle (Saale). Halle (Saale).
- [Nelis et al. 2010] ^ Nelis, Mari, Tõnu Esko, Reedik Mägi, Fritz Zimprich, Alexander Zimprich, Draga Toncheva, Sena Karachanak, Tereza Piskáčková, Ivan Balaščák, Leena Peltonen, Eveliina Jakkula, Karola Rehnström, Mark Lathrop, Simon Heath, Pilar Galan, Stefan Schreiber, Thomas Meitinger, Arne Pfeufer, H. Erich Wichmann, Béla Melegh, Noémi Polgár, Daniela Toniolo, Paolo Gasparini, Pio D'Adamo, Janis Klovins, Liene Nikitina-Zake, Vaidutis Kučinskas, Jūratė Kasnauskienė, Jan Lubinski, Tadeusz Debniak, Svetlana Limborska, Andrey Khrunin, Xavier Estivill, Raquel Rabionet, Sara Marsal, Antonio Julià, Stylianos E. Antonarakis, Samuel Deutsch, Christelle Borel, Homa Attar, Maryline Gagnebin, Milan Macek, Michael Krawczak, Maido Remm, and Andres Metspalu. 2010. Correction: Genetic Structure of Europeans: A View from the North–East. PLOS ONE 5 (3) https://doi.org/10.1371/annotation/2849e182-aef5-4e2b-a5ac-0b74b30e5f48
- [Översti et al. 2017] ^ Översti, Sanni, Päivi Onkamo, Monika Stoljarova, Bruce Budowle, Antti Sajantila, and Jukka U. Palo. 2017. Identification and analysis of mtDNA genomes attributed to Finns reveal long-stagnant demographic trends obscured in the total diversity. Scientific Reports 7 (1):6193. https://doi.org/10.1038/s41598-017-05673-7
- [Rosser et al. 2000] ^ Rosser, Zoë H., Tatiana Zerjal, Matthew E. Hurles, Maarja Adojaan, Dragan Alavantic, António Amorim, William Amos, Manuel Armenteros, Eduardo Arroyo, Guido Barbujani, Gunhild Beckman, Lars Beckman, Jaume Bertranpetit, Elena Bosch, Daniel G. Bradley, Gaute Brede, Gillian Cooper, Helena B. S. M. Côrte-Real, Peter de Knijff, Ronny Decorte, Yuri E. Dubrova, Oleg Evgrafov, Anja Gilissen, Sanja Glisic, Mukaddes Gölge, Emmeline W. Hill, Anna Jeziorowska, Luba Kalaydjieva, Manfred Kayser, Toomas Kivisild, Sergey A. Kravchenko, Astrida Krumina, Vaidutis Kučinskas, João Lavinha, Ludmila A. Livshits, Patrizia Malaspina, Syrrou Maria, Ken McElreavey, Thomas A. Meitinger, Aavo-Valdur Mikelsaar, R. John Mitchell, Khedoudja Nafa, Jayne Nicholson, Søren Nørby, Arpita Pandya, Jüri Parik, Philippos C. Patsalis, Luísa Pereira, Borut Peterlin, Gerli Pielberg, Maria João Prata, Carlo Previderé, Lutz Roewer, Siiri Rootsi, D. C. Rubinsztein, Juliette Saillard, Fabrício R. Santos, Gheorghe Stefanescu, Bryan C. Sykes, Aslihan Tolun, Richard Villems, Chris Tyler-Smith, and Mark A. Jobling. 2000. Y-Chromosomal Diversity in Europe Is Clinal and Influenced Primarily by Geography, Rather than by Language. The American Journal of Human Genetics 67 (6):1526-1543.
- [Tambets et al. 2004] ^ Tambets, Kristiina, Siiri Rootsi, Toomas Kivisild, Hela Help, Piia Serk, Eva-Liis Loogväli, Helle-Viivi Tolk, Maere Reidla, Ene Metspalu, Liana Pliss, Oleg Balanovsky, Andrey Pshenichnov, Elena Balanovska, Marina Gubina, Sergey Zhadanov, Ludmila Osipova, Larisa Damba, Mikhail Voevoda, Ildus Kutuev, Marina Bermisheva, Elza Khusnutdinova, Vladislava Gusar, Elena Grechanina, Jüri Parik, Erwan Pennarun, Christelle Richard, Andre Chaventre, Jean-Paul Moisan, Lovorka Barać, Marijana Peričić, Pavao Rudan, Rifat Terzić, Ilia Mikerezi, Astrida Krumina, Viesturs Baumanis, Slawomir Koziel, Olga Rickards, Gian Franco De Stefano, Nicholas Anagnou, Kalliopi I. Pappa, Emmanuel Michalodimitrakis, Vladimir Ferák, Sandor Füredi, Radovan Komel, Lars Beckman, and Richard Villems. 2004. The Western and Eastern Roots of the Saami—the Story of Genetic “Outliers” Told by Mitochondrial DNA and Y Chromosomes. The American Journal of Human Genetics 74 (4):661-682. http://dx.doi.org/10.1086/383203