Corded Ware migration


The Corded Ware culture has been argued to practice exogamy – most adult women being of non-local origin – based on a recent work on diet and mobility[Sjogren, Price, and Kristiansen 2016], and mtDNA has been documented to be more varied among Corded Ware females than men[Lazaridis et al. 2014]. Exogamy is described as a well-established practice over a long period of time, and the origin of females has been linked to Neolithic cultures still residing on the higher elevations in the region shared with Corded Ware cultures, which colonised rivers and valleys[Kristiansen et al. 2017].

A violent picture has been proposed for the Corded Ware culture society[Haak et al. 2008], with warrior youth bands with seasonal activities, in a way similar to that documented in the Russian steppe from the Bronze Age onwards[Kristiansen et al. 2017].

In a first stage of the CWC migrations, their nomadic way of life is represented by site-camps close to tumuli, located on routes of their long-lasting travels. In a second phase, semi-nomadic economic allowed for settlement micro-regions to appear, and cemeteries were a kind of landmark during the population’s seasonal migrations[Machnik 2004]. Necropolises were significant areas to which they returned periodically, establishing eventually site-camps, beginning their transformation into Early Bronze Age groups with a more stable economy[Witkowska 2006].

Analysis of aDNA has revealed that the plague was a prehistoric disease, associated with the Eurasian steppes, and linked to the Corded Ware culture expansion[Rasmussen et al. 2015][Andrades Valtueña et al. 2017], which connected vast areas in central-east Europe in a relatively short period. Also, the population to the north of the loess belt of the northern European plain is known to have been much lesser than in south-east Europe[Müller 2013].

This demographic disadvantage of central-east European lands might have provided part of the ‘push’ for the migration and expansion of the Corded Ware population[Anthony and Brown 2017], which show a clear genetic homogeneity – close to the so-called ‘Yamna component’ – from western to eastern Europe, in spite of the proposed practice of exogamy. This explosive migration over a sparsely populated area accounts for the expansion of R1a1a1b-Z645 lineages, possibly originally from the Dniester-Dnieper or Middle Dnieper region.

The most recent sample from Zvejnieki, of the Late Neolithic / Corded Ware (ca. 2885 BC), clusters quite closely with Yamna samples, revealing a break of the regional population with the samples from a thousand years earlier, which were closer to EHG[Mathieson et al. 2017]. The lack of more EHG ancestry – as expected in a sample potentially generations away from southern migrants (and thus admixed with the local population), and the elevated steppe component found – closer to Yamna samples than most Sredni Stog and Corded Ware samples – point to a recent contribution from the steppe, potentially through exogamy.

Its mtDNA haplogroup is U5a1b. Although different U5, U4, and U2 subclades are commonly found in hunter-gatherer groups, this precise haplogroup is found previously only in four Sredni Stog samples from Deriivka (ca. 5150 BC), and later in central European Benzingerode (ca. 2275 BC), and in a sample from Przecławice (ca. 1790 BC), of the Únětice culture[Haak et al. 2015][Mathieson et al. 2017]. This may give support to the origin of the Corded Ware culture near the region of the Sredni Stog culture, and points to the north-west Pontic population (and women in particular) as a potential origin of the Yamna ancestry found in the sample.

european-hunter-gatherer.jpg Image modified from Mathieson et al. (2017). «Structure and population change in European populations with mainly hunter gatherer ancestry. Background: Effective migration surface inferred using 109 Mesolithic and Early Neolithic genomes. Blue indicates regions of high migration and red regions of low migration (relative migration rate m). Pies: Each pie represents one or more individuals. Blue, red and green segments pies show, respectively, WHG, EHG and CHG ancestry – we chose a 2-population model if it fitted in qpAdm with p>0.01. Pies with 100% ancestry are those that were fixed as source populations. In Latvia, Samara and Ukraine, sets of three pies show the transition in ancestries, approximately from Mesolithic- Neolithic-Yamnaya. Populations with labels in brackets were not used to fit the migration surface. B: Ancestry transitions for hunter-gatherer populations in the 3-population WHG+EHG+CHG model. Dashed lines show populations with geographic continuity». Original under a CC-BY-NC 4.0 International license.

R1a1a1b-Z645, split from R1a1a1-M417 ca. 3500 BC, shows a TMRCA of ca. 3000 BC, coinciding with the formation date for mainly-European subclades R1a1a1b1-Z283 and R1a1a1b1a-Z282, and mainly-Asian subclade R1a1a1b2-Z93. The common TMRCA for R1a1a1b1-Z283 and R1a1a1b1a-Z282 suggests an expansion at nearly the same time as peoples of Corded Ware cultures are supposed to have migrated east- and westward, reaching the Middle Elbe-Saale region – where most aDNA samples analysed come from – about 2750 BC. The common TMRCA of 2700 BC for modern Asian lineages gives support to a later successful expansion into Asia centred on the eastern part of the Pontic-Caspian steppes (see Indo-Iranian).

Y-DNA samples of haplogroup R1a-M420 (probably R1a1a1-M417) are found in central Corded Ware culture groups[Allentoft et al. 2015][Mathieson et al. 2015][Haak et al. 2008][Mittnik et al. 2017][Saag et al. 2017]: one sample in Bergrheinfeld (ca. 2647), two in Eulau (ca. 2600 BC), one[1] from Kyndelöse (ca. 2670 BC), seven from Esperstedt (one dated ca. 2430 BC and other six ca. 2275 BC), and two from the Battle Axe culture – one from Viby (ca. 2500 BC), and one from Ölsund (ca. 2350 BC).

Five samples from Esperstedt have been inferred to be relatives via paternal line, which confirms their common sharing of bottom subclade R1a1a1-M417[Monroy Kuhn, Jakobsson, and Günther 2017]. The oldest sample from Esperstedt (labelled I0104), a second-degree relative to the rest, has been found to cluster the closest to steppe samples, closer than any other Corded Ware sample, previous or posterior, or any samples from Sintashta, or Potapovka. This, connected with the exogamy prevalent among Corded Ware peoples, and the nomadic nature of its culture, precludes a proper interpretation of the ancestry found in the family, since it might have come quite recently directly from the steppe, at a time when the Catacomb and Potapovka cultures had already replaced the Yamna culture.

In terms of mtDNA the Corded Ware culture shows common hunter-gatherer haplogroups U5, U2, or U4 (Brandt et al. 2013). Out of seventeen samples from the Corded Ware culture, four (all of Esperstedt), include different J1c subclades, only found previously accompanying the expansion of Neolithic Middle Eastern farmers, including the Globular Amphora culture (in two out of nine samples), and later in one out of five samples of the Sintashta culture[Mathieson et al. 2017].


Image modified from Monroy Kuhn, Jakobsson, and Günther (2017). «Kin-relationship among males at the Corded Ware site in Esperstedt, Germany. (A) The five individuals, their inferred degree of relationship and their uniparental haplogroups. The dashed line between I1540 and I1538 shows a second degree relationship missed by READ. (B) Map of the Corded Ware site (reference site 4) near Esperstedt, Germany. Blue circles show the locations of Corded Ware Burials. The approximate burial locations of the individuals with inferred relationships are indicated by their ID. Map data © OpenStreetMap contributors, CC BY-SA.» Image under a CC-BY-NC 4.0 International license.

Two samples from Tiefbrunn of haplogroups R-M207 (ca. 2755 BC) and R1b1-L278 (ca. 2725 BC) are of dubious subclade – the first could be R1a1a1-M417 or R1b1-L278, the second might correspond to older European hunter-gatherer lineages, or they might be associated with the expansion of Corded Ware cultures from the Balkans. Other non-R-M207 samples from central Corded Ware cultures include the two oldest ones, of IJ and G2a-P15 lineages, from Jagodno ca. 2800 BC[Gworys et al. 2013]; and one of haplogroup P-P295 in Esperstedt from ca. 2275 BC.

In the Baltic and the Forest zone, four samples are dated around 2500 BC: haplogroups R1a1a1b-Z645 and R1a1a1b1-Z283 in Kunila, and two samples of haplogroup R1a1a1b-Z645 in Ardu. These samples together with its previous presence in Usvyatyan culture (ca. 2500 BC) and in Naumovo and Sertaya II[Chekunova et al. 2014], and its continuity in later times suggest that R1a1a1b-Z645 lineages almost fully replaced the previous R1b-M343 lineages in the eastern Baltic around the time of the Corded Ware culture expansion.

plot5_cut_CW.png Detail of PCA analysis of free datasets including Minoans and Mycenaeans[Lazaridis et al. 2017], and Scythian and Sarmatian[Unterländer et al. 2017] samples. PC2 vs. PC1. The graphic has been arranged so that ancestries and samples are located in geographically friendly axes similar to north-south (Y), east-west(X). Symbols are used, in a simplified manner, in accordance with symbols for Y-DNA haplogroups used in the maps. Labels have been used for simplification. Areas are drawn surrounding Yamna/Poltavka, Corded Ware (including samples from Estonia, Battle Axe, and Poltavka outlier), and succeeding Sintashta and Andronovo cultures, as well as Bell Beaker. Corded Ware sample I0104, from Erperstedt, has also been labelled.

The oldest R1a1a1-M417 lineages of Central Europe are found in early Corded Ware groups, while ancient DNA from Neolithic Linear Pottery (ca. 5500–3500 BC) and Globular Amphora (ca. 3400-2800 BC) cultures have been found to correspond mainly to I2-M438 and G2a-P15 lineages, with no steppe-related ancestry in admixture analyses[Haak et al. 2015][Mathieson et al. 2017]. This raises the possibility of a wide and rapid spread of R1a1a1-M417, and especially R1a1a1b1-Z283 subclades in Europe[Underhill et al. 2015] associated with the expansion of the Corded Ware culture. The diversification of R1a1a1b2-Z93 in the Middle East and South Asia remains more obscure[Underhill et al. 2015].

However, in samples obtained from Estonia it was seen that admixture between Corded Ware culture farmers and Comb Ceramic culture hunter-gatherers may have been limited in males of R1a1a1-M417 lineages. Also, the presence of a genetic component associated with Caucasus hunter-gatherers – also present in Yamna migrants, Eastern hunter-gatherers, and individuals from the Estonian Comb Ceramic culture, means that the expansion of the Corded Ware culture cannot be seen as the sole means for the spread of this genetic component, at least in eastern Europe[Saag et al. 2017].

copper-age-early_cut.jpg Diachronic map of Copper Age migrations ca. 3100-2600 BC [Anthony 2007][Harrison and Heyd 2007][Sjogren, Price, and Kristiansen 2016][Heyd 2012][Heyd 2014].


  • [Allentoft et al. 2015] ^ Allentoft, Morten E., Martin Sikora, Karl-Goran Sjogren, Simon Rasmussen, Morten Rasmussen, Jesper Stenderup, Peter B. Damgaard, Hannes Schroeder, Torbjorn Ahlstrom, Lasse Vinner, Anna-Sapfo Malaspinas, Ashot Margaryan, Tom Higham, David Chivall, Niels Lynnerup, Lise Harvig, Justyna Baron, Philippe Della Casa, Pawel Dabrowski, Paul R. Duffy, Alexander V. Ebel, Andrey Epimakhov, Karin Frei, Miroslaw Furmanek, Tomasz Gralak, Andrey Gromov, Stanislaw Gronkiewicz, Gisela Grupe, Tamas Hajdu, Radoslaw Jarysz, Valeri Khartanovich, Alexandr Khokhlov, Viktoria Kiss, Jan Kolar, Aivar Kriiska, Irena Lasak, Cristina Longhi, George McGlynn, Algimantas Merkevicius, Inga Merkyte, Mait Metspalu, Ruzan Mkrtchyan, Vyacheslav Moiseyev, Laszlo Paja, Gyorgy Palfi, Dalia Pokutta, Lukasz Pospieszny, T. Douglas Price, Lehti Saag, Mikhail Sablin, Natalia Shishlina, Vaclav Smrcka, Vasilii I. Soenov, Vajk Szeverenyi, Gusztav Toth, Synaru V. Trifanova, Liivi Varul, Magdolna Vicze, Levon Yepiskoposyan, Vladislav Zhitenev, Ludovic Orlando, Thomas Sicheritz-Ponten, Soren Brunak, Rasmus Nielsen, Kristian Kristiansen, and Eske Willerslev. 2015. Population genomics of Bronze Age Eurasia. Nature 522 (7555):167-172.
  • [Andrades Valtueña et al. 2017] ^ Andrades Valtueña, Aida, Alissa Mittnik, Felix M. Key, Wolfgang Haak, Raili Allmäe, Andrej Belinskij, Mantas Daubaras, Michal Feldman, Rimantas Jankauskas, Ivor Janković, Ken Massy, Mario Novak, Saskia Pfrengle, Sabine Reinhold, Mario Šlaus, Maria A. Spyrou, Anna Szecsenyi-Nagy, Mari Tõrv, Svend Hansen, Kirsten I. Bos, Philipp W. Stockhammer, Alexander Herbig, and Johannes Krause. 2017. The Stone Age Plague: 1000 years of Persistence in Eurasia. bioRxiv.
  • [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.
  • [Anthony and Brown 2017] ^ Anthony, D.W., and D. R. Brown. 2017. Molecular Archaeology and Indo-European linguistics: Impressions from new data. In Usque ad Radices: Indo-European Studies in Honour of Birgit Anette Olsen, edited by B. Simmelkjær, S. Hansen, A. Hyllested, A. R. Jørgensen, G. Kroonen, J. H. Larsson, B. N. Whitehead, T. Olander and T. M. Søborg. Copenhagen: Museum Tusculanum 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.
  • [Gworys et al. 2013] ^ Gworys, Bohdan, Joanna Rosińczuk-Tonderys, Aleksander Chrószcz, Maciej Janeczek, Andrzej Dwojak, Justyna Bazan, Mirosław Furmanek, Tadeusz Dobosz, Małgorzata Bonar, Anna Jonkisz, and Ireneusz Całkosiński. 2013. Assessment of late Neolithic pastoralist's life conditions from the Wroclaw–Jagodno site (SW Poland) on the basis of physiological stress markers. Journal of Archaeological Science 40 (6):2621-2630.
  • [Haak et al. 2008] ^ 1 2 Haak, W., G. Brandt, H. N. de Jong, C. Meyer, R. Ganslmeier, V. Heyd, C. Hawkesworth, A. W. Pike, H. Meller, and K. W. Alt. 2008. Ancient DNA, Strontium isotopes, and osteological analyses shed light on social and kinship organization of the Later Stone Age. Proc Natl Acad Sci U S A 105 (47):18226-31.
  • [Haak et al. 2015] ^ 1 2 Haak, W., I. Lazaridis, N. Patterson, N. Rohland, S. Mallick, B. Llamas, G. Brandt, S. Nordenfelt, E. Harney, K. Stewardson, Q. Fu, A. Mittnik, E. Banffy, C. Economou, M. Francken, S. Friederich, R. G. Pena, F. Hallgren, V. Khartanovich, A. Khokhlov, M. Kunst, P. Kuznetsov, H. Meller, O. Mochalov, V. Moiseyev, N. Nicklisch, S. L. Pichler, R. Risch, M. A. Rojo Guerra, C. Roth, A. Szecsenyi-Nagy, J. Wahl, M. Meyer, J. Krause, D. Brown, D. Anthony, A. Cooper, K. W. Alt, and D. Reich. 2015. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522 (7555):207-11.
  • [Harrison and Heyd 2007] ^ Harrison, Richard, and Volker Heyd. 2007. The Transformation of Europe in the Third Millennium BC: the example of ‘Le Petit-Chasseur I + III’ (Sion, Valais, Switzerland). Praehistorische Zeitschrift 82 (2).
  • [Heyd 2012] ^ Heyd, Volker. 2012. Yamnaya gropus and tumuli west of the Black Sea. Travaux de la Maison de l'Orient et de la Méditerranée. Série recherches archéologiques 58 (1):535-555.
  • [Heyd 2014] ^ Heyd, Volker. 2014. Families, Prestige Goods, Warriors & Complex Societies: Beaker Groups of the 3rd Millennium cal BC Along the Upper & Middle Danube. Proceedings of the Prehistoric Society 73:327-379.
  • [Kristiansen et al. 2017] ^ 1 2 Kristiansen, Kristian, Morten E. Allentoft, Karin M. Frei, Rune Iversen, Niels N. Johannsen, Guus Kroonen, Łukasz Pospieszny, T. Douglas Price, Simon Rasmussen, Karl-Göran Sjögren, Martin Sikora, and Eske Willerslev. 2017. Re-theorising mobility and the formation of culture and language among the Corded Ware Culture in Europe. Antiquity 91 (356):334-347.
  • [Lazaridis et al. 2014] ^ Lazaridis, I., N. Patterson, A. Mittnik, G. Renaud, S. Mallick, K. Kirsanow, P. H. Sudmant, J. G. Schraiber, S. Castellano, M. Lipson, B. Berger, C. Economou, R. Bollongino, Q. Fu, K. I. Bos, S. Nordenfelt, H. Li, C. de Filippo, K. Prufer, S. Sawyer, C. Posth, W. Haak, F. Hallgren, E. Fornander, N. Rohland, D. Delsate, M. Francken, J. M. Guinet, J. Wahl, G. Ayodo, H. A. Babiker, G. Bailliet, E. Balanovska, O. Balanovsky, R. Barrantes, G. Bedoya, H. Ben-Ami, J. Bene, F. Berrada, C. M. Bravi, F. Brisighelli, G. B. Busby, F. Cali, M. Churnosov, D. E. Cole, D. Corach, L. Damba, G. van Driem, S. Dryomov, J. M. Dugoujon, S. A. Fedorova, I. Gallego Romero, M. Gubina, M. Hammer, B. M. Henn, T. Hervig, U. Hodoglugil, A. R. Jha, S. Karachanak-Yankova, R. Khusainova, E. Khusnutdinova, R. Kittles, T. Kivisild, W. Klitz, V. Kucinskas, A. Kushniarevich, L. Laredj, S. Litvinov, T. Loukidis, R. W. Mahley, B. Melegh, E. Metspalu, J. Molina, J. Mountain, K. Nakkalajarvi, D. Nesheva, T. Nyambo, L. Osipova, J. Parik, F. Platonov, O. Posukh, V. Romano, F. Rothhammer, I. Rudan, R. Ruizbakiev, H. Sahakyan, A. Sajantila, A. Salas, E. B. Starikovskaya, A. Tarekegn, D. Toncheva, S. Turdikulova, I. Uktveryte, O. Utevska, R. Vasquez, M. Villena, M. Voevoda, C. A. Winkler, L. Yepiskoposyan, P. Zalloua, T. Zemunik, A. Cooper, C. Capelli, M. G. Thomas, A. Ruiz-Linares, S. A. Tishkoff, L. Singh, K. Thangaraj, R. Villems, D. Comas, R. Sukernik, M. Metspalu, M. Meyer, E. E. Eichler, J. Burger, M. Slatkin, S. Paabo, J. Kelso, D. Reich, and J. Krause. 2014. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513 (7518):409-13.
  • [Lazaridis et al. 2017] ^ Lazaridis, Iosif, Alissa Mittnik, Nick Patterson, Swapan Mallick, Nadin Rohland, Saskia Pfrengle, Anja Furtwängler, Alexander Peltzer, Cosimo Posth, Andonis Vasilakis, P. J. P. McGeorge, Eleni Konsolaki-Yannopoulou, George Korres, Holley Martlew, Manolis Michalodimitrakis, Mehmet Özsait, Nesrin Özsait, Anastasia Papathanasiou, Michael Richards, Songül Alpaslan Roodenberg, Yannis Tzedakis, Robert Arnott, Daniel M. Fernandes, Jeffery R. Hughey, Dimitra M. Lotakis, Patrick A. Navas, Yannis Maniatis, John A. Stamatoyannopoulos, Kristin Stewardson, Philipp Stockhammer, Ron Pinhasi, David Reich, Johannes Krause, and George Stamatoyannopoulos. 2017. Genetic origins of the Minoans and Mycenaeans. Nature 548 (7666):214-218.
  • [Machnik 2004] ^ Machnik, J. 2004. Pasterstwo u społeczeństw kultury ceramiki sznurowej w dorzeczu górnej Wisły, górnego Bugu i Dniestru. In Nomadyzm a pastoralizm w mię-dzyrzeczu Wisły i Dniepru (neolit, eneolit, epoka brązu),, edited by A. Kośko and M. Szmyt. Poznań.
  • [Mathieson et al. 2015] ^ Mathieson, I., I. Lazaridis, N. Rohland, S. Mallick, N. Patterson, S. A. Roodenberg, E. Harney, K. Stewardson, D. Fernandes, M. Novak, K. Sirak, C. Gamba, E. R. Jones, B. Llamas, S. Dryomov, J. Pickrell, J. L. Arsuaga, J. M. de Castro, E. Carbonell, F. Gerritsen, A. Khokhlov, P. Kuznetsov, M. Lozano, H. Meller, O. Mochalov, V. Moiseyev, M. A. Guerra, J. Roodenberg, J. M. Verges, J. Krause, A. Cooper, K. W. Alt, D. Brown, D. Anthony, C. Lalueza-Fox, W. Haak, R. Pinhasi, and D. Reich. 2015. Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528 (7583):499-503.
  • [Mathieson et al. 2017] ^ 1 2 3 4 Mathieson, Iain, Songül Alpaslan Roodenberg, Cosimo Posth, Anna Szécsényi-Nagy, Nadin Rohland, Swapan Mallick, Iñigo Olade, Nasreen Broomandkhoshbacht, Olivia Cheronet, Daniel Fernandes, Matthew Ferry, Beatriz Gamarra, Gloria González Fortes, Wolfgang Haak, Eadaoin Harney, Ben Krause-Kyora, Isil Kucukkalipci, Megan Michel, Alissa Mittnik, Kathrin Nägele, Mario Novak, Jonas Oppenheimer, Nick Patterson, Saskia Pfrengle, Kendra Sirak, Kristin Stewardson, Stefania Vai, Stefan Alexandrov, Kurt W. Alt, Radian Andreescu, Dragana Antonović, Abigail Ash, Nadezhda Atanassova, Krum Bacvarov, Mende Balázs Gusztáv, Hervé Bocherens, Michael Bolus, Adina Boroneanţ, Yavor Boyadzhiev, Alicja Budnik, Josip Burmaz, Stefan Chohadzhiev, Nicholas J. Conard, Richard Cottiaux, Maja Čuka, Christophe Cupillard, Dorothée G. Drucker, Nedko Elenski, Michael Francken, Borislava Galabova, Georgi Ganetovski, Bernard Gely, Tamás Hajdu, Veneta Handzhyiska, Katerina Harvati, Thomas Higham, Stanislav Iliev, Ivor Janković, Ivor Karavanić, Douglas J. Kennett, Darko Komšo, Alexandra Kozak, Damian Labuda, Martina Lari, Catalin Lazar, Maleen Leppek, Krassimir Leshtakov, Domenico Lo Vetro, Dženi Los, Ivaylo Lozanov, Maria Malina, Fabio Martini, Kath McSweeney, Harald Meller, Marko Menđušić, Pavel Mirea, Vyacheslav Moiseyev, Vanya Petrova, T. Douglas Price, Angela Simalcsik, Luca Sineo, Mario Šlaus, Vladimir Slavchev, Petar Stanev, Andrej Starović, Tamás Szeniczey, Sahra Talamo, Maria Teschler-Nicola, Corinne Thevenet, Ivan Valchev, Frédérique Valentin, Sergey Vasilyev, Fanica Veljanovska, Svetlana Venelinova, Elizaveta Veselovskaya, Bence Viola, Cristian Virag, Joško Zaninović, Steve Zäuner, Philipp W. Stockhammer, Giulio Catalano, Raiko Krauß, David Caramelli, Gunita Zariņa, Bisserka Gaydarska, Malcolm Lillie, Alexey G. Nikitin, Inna Potekhina, Anastasia Papathanasiou, Dušan Borić, Clive Bonsall, Johannes Krause, Ron Pinhasi, and David Reich. 2017. The Genomic History Of Southeastern Europe. bioRxiv.
  • [Mittnik et al. 2017] ^ Mittnik, Alissa, Chuan-Chao Wang, Saskia Pfrengle, Mantas Daubaras, Gunita Zariņa, Fredrik Hallgren, Raili Allmäe, Valery Khartanovich, Vyacheslav Moiseyev, Anja Furtwängler, Aida Andrades Valtueña, Michal Feldman, Christos Economou, Markku Oinonen, Andrejs Vasks, Mari Tõrv, Oleg Balanovsky, David Reich, Rimantas Jankauskas, Wolfgang Haak, Stephan Schiffels, and Johannes Krause. 2017. The Genetic History of Northern Europe. bioRxiv.
  • [Müller 2013] ^ Müller, J. 2013. Demographic traces of technological innovation, social change and mobility: from 1 to 8 million Europeans (6000-2000 BCE). In Environment and subsistence – forty years after Janusz Kruk’s „Settlement studies…” (= Studien zur Archäologie in Ostmitteleuropa / Studia nad Pradziejami Europy Środkowej 11), edited by S. Kadrow and P. Włodarczak. Rzeszów, Bonn: Mitel & Verlag Dr. Rudolf Habelt.
  • [Rasmussen et al. 2015] ^ Rasmussen, Simon, Morten Erik Allentoft, Kasper Nielsen, Ludovic Orlando, Martin Sikora, Karl-Göran Sjögren, Anders Gorm Pedersen, Mikkel Schubert, Alex Van Dam, Christian Moliin Outzen Kapel, Henrik Bjørn Nielsen, Søren Brunak, Pavel Avetisyan, Andrey Epimakhov, Mikhail Viktorovich Khalyapin, Artak Gnuni, Aivar Kriiska, Irena Lasak, Mait Metspalu, Vyacheslav Moiseyev, Andrei Gromov, Dalia Pokutta, Lehti Saag, Liivi Varul, Levon Yepiskoposyan, Thomas Sicheritz-Pontén, Robert A Foley, Marta Mirazón Lahr, Rasmus Nielsen, Kristian Kristiansen, and Eske Willerslev. 2015. Early Divergent Strains of Yersinia pestis in Eurasia 5,000 Years Ago. Cell 163 (3):571-582.
  • [Saag et al. 2017] ^ 1 2 Saag, Lehti, Liivi Varul, Christiana Lyn Scheib, Jesper Stenderup, Morten E Allentoft, Lauri Saag, Luca Pagani, Maere Reidla, Kristiina Tambets, Ene Metspalu, Aivar Kriiska, Eske Willerslev, Toomas Kivisild, and Mait Metspalu. 2017. Extensive farming in Estonia started through a sex-biased migration from the Steppe. bioRxiv.
  • [Szmyt 2013] Szmyt, Marzena. 2013. The circulation of People and Ideas in the Baltic and Pontic Areas during 3rd millennium BC.
  • [Underhill et al. 2015] ^ 1 2 Underhill, P. A., G. D. Poznik, S. Rootsi, M. Jarve, A. A. Lin, J. Wang, B. Passarelli, J. Kanbar, N. M. Myres, R. J. King, J. Di Cristofaro, H. Sahakyan, D. M. Behar, A. Kushniarevich, J. Sarac, T. Saric, P. Rudan, A. K. Pathak, G. Chaubey, V. Grugni, O. Semino, L. Yepiskoposyan, A. Bahmanimehr, S. Farjadian, O. Balanovsky, E. K. Khusnutdinova, R. J. Herrera, J. Chiaroni, C. D. Bustamante, S. R. Quake, T. Kivisild, and R. Villems. 2015. The phylogenetic and geographic structure of Y-chromosome haplogroup R1a. Eur J Hum Genet 23 (1):124-31.
  • [Unterländer et al. 2017] ^ Unterländer, Martina, Friso Palstra, Iosif Lazaridis, Aleksandr Pilipenko, Zuzana Hofmanová, Melanie Groß, Christian Sell, Jens Blöcher, Karola Kirsanow, Nadin Rohland, Benjamin Rieger, Elke Kaiser, Wolfram Schier, Dimitri Pozdniakov, Aleksandr Khokhlov, Myriam Georges, Sandra Wilde, Adam Powell, Evelyne Heyer, Mathias Currat, David Reich, Zainolla Samashev, Hermann Parzinger, Vyacheslav I. Molodin, and Joachim Burger. 2017. Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe. 8:14615.
  • [Witkowska 2006] ^ Witkowska, Barbara. 2006. Corded Ware Culture Settlements on Central European Uplands. Sprawozdania Archeologiczne 58:21-100.


  1. Published as corresponding to Nordic Middle Neolithic culture, additional information by Vladimir Tagankin revealed a branch typical of modern Nordic (R1a1a1b1a3-Z284) subclades, and a new date of ca. 2475 BC, including reduction for high marine signal.