A more recent, revised and updated version of this paper has been published (2019)

Bell Beaker expansion

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Steppe ancestry has been recently found widespread in central European Bell Beaker individuals, while in Iberian samples this ancestry was not found, therefore excluding a substantial contribution to central Europe from Iberian Beaker Complex-associated individuals[Olalde et al. 2017]. This further supports Heyd’s archaeological model[Harrison and Heyd 2007][Heyd 2012][Heyd 2014], and contradicts the previous model of population expansion out of Iberia.


Modified file from recent papers on ancient samples from Eastern European, Southeastern European, Western European, and Bell Beaker cultures: Left: ADMIXTURE clustering analysis with k=8 showing ancient individuals. E/M/MLN, Early/Middle/Middle Late Neolithic; W/E/S/CHG, Western/Eastern/Scandinavian/Caucasus hunter-gatherers[Olalde et al. 2017]. Center: Supervised ADMIXTURE plot, modeling each ancient individual (one per row), as a mixture of populations represented by clusters containing Anatolian Neolithic (grey), Yamnaya from Samara (yellow), EHG (pink) and WHG (green). Dates indicate approximate range of individuals in each population[Mathieson et al. 2017]. Right: Ancestral components in ancient individuals estimated by ADMIXTURE (k=11)[Mittnik et al. 2017]. Original images under a CC-BY-NC 4.0 International license.

In the lower Danube clear patterns of patrilocality and female exogamy have been found, apart from a continuing kinship relation in the transition of the Chalcolithic to the Bronze Age. There is evidence of continuing traditions from the Bell Beaker cultures to Early Bronze Age cultures in the region, with female mobility as a force for regional and supraregional communication and exchange[Knipper et al. 2017].


Image from Olalde et al. (2017). «Proportion of Steppe-related ancestry (shown in black) in Beaker Complex groups, computed with qpAdm under the model Yamnaya_Samara + Anatolia_Neolithic + WHG. The area of the pie is proportional to the number of individuals (shown inside the pie if more than one)»”. Original images under a CC-BY-NC 4.0 International license.

Bell Beaker samples from Hungary at overlapping dates show very different proportions (from 0% to 74%) of steppe ancestry, a heterogeneity consistent with early stages of admixture between European farmers and Yamna migrants. Also, in southern France and Britain there is a greater affinity to the Iberian Early Neolithic farmers than to central European farmers, which suggests a previous spread of migrants along the Atlantic coast, consistent with the interaction of Megalithic-building cultures involving movements of people[Olalde et al. 2017].

Bell Beaker samples from Hungary at overlapping dates show very different proportions (from 0% to 74%) of steppe ancestry, a heterogeneity consistent with early stages of admixture between European farmers and Yamna migrants. Also, in southern France and Britain there is a greater affinity to the Iberian Early Neolithic farmers than to central European farmers, which suggests a previous spread of migrants along the Atlantic coast, consistent with the interaction of Megalithic-building cultures involving movements of people (Olalde et al. 2017).


Image from Olalde et al. (2017). «Principal component analysis of 990 present-day West Eurasian individuals (grey dots), with previously published (pale yellow) and new ancient samples projected onto the first two principal components»”. Original images under a CC-BY-NC 4.0 International license.

The oldest samples from the Bell Beaker culture are two individuals from Kromsdorf dated ca. 2550 BC, one of R1b1a1a2-M269 (xR1b1a1a2a1a1-U106), and the other of R1b-M343 (M269 unclear) lineages[Lee et al. 2012]. The oldest samples of R1b1a1a2a1a2-P312 lineages are found in Osterhofen ca. 2540 BC, and in Sierentz – Les villas d’Aurèle ca. 2430 BC. The oldest sample of haplogroup R1b1a1a2a1a2b-U152 is found in Budapest ca. 2335 BC, of haplogroup R1b1a1a2a1a2a-DF27[1] in Quedlinburg ca. 2290 BC[Lazaridis et al. 2016], and of haplogroup R1b1a1a2a1a2c-L21 in Amesbury ca. 2290 BC[Olalde et al. 2017].

chalcolithic_late_Europe.jpg Diachronic map of migrations in Europe ca. 2600-2250 BC[Heyd 2014][Anthony 2007][Sjogren, Price, and Kristiansen 2016][Harrison and Heyd 2007][Prieto Martínez and Salanova 2015][Fokkens and Nicolis 2012].

Regarding the arrival of Bell Beakers into Britain – marked by the appearance of steppe ancestry in aDNA samples –, it has been found that they are more closely related to lower Rhine individuals, and not to the Atlantic façade of western Europe[Olalde et al. 2017]. Demic diffusion of R1b1a1a2a1a2c1-L21 lineages accompanying Bell Beaker expansion in the British Isles was already supported by ancient DNA analysis[Cassidy et al. 2016], and all samples of the British Isles have been reported as of R1b1a1a2a1a2c-L21 (or older haplogroups). These data and the modern distribution of L21 subclades associated with the British Isles point more likely to a single successful migration of clans of R1b1a1a2a1a2c-L21 lineages into Britain. The contended Pre-Celtic Irish and Pictish substrates might have been therefore of Late Indo-European nature, imported by peoples of R1b1a1a2a1a2c1-L21 lineages.

Recent research already supported a considerable degree of mobility with little difference between male and female migration in Britain[Parker Pearson et al. 2016], and not an exchange of female marriage partners[Brodie 2001] or inter-cultural contact consolidation[Vander Linden 2007], as previously proposed.

r1b-L21.jpg Modern distribution of haplogroup R1b1a1a2a1a2c1-L21. Adapted from Myres et al.[Myres et al. 2011]

Studies of ancient Indo-European hydronymy[Krahe 1964][Krahe 1949][Nicolaisen 1957] have revealed a quasi-uniform name-giving system for water courses that shows Indo-European water-words and suffixes following rules of Late Proto-Indo-European word formation[Adrados 1998]. This points to an ancient wave of Late Indo-European speakers spread over Western and Central Europe before the Celtic and Germanic expansions, including the British Isles, the Italian and Iberian peninsulas.

The four certain samples of R1b1a1a2a1a2b-U152 lineage have been found in Bell Beaker territories from east to west Europe. The presence of a potential R1b1a1a2a1a2a-DF27 lineage in the central Bell Beaker group, and its subclade distribution in the modern west European population might be linked to this lineage’s expansion to the west and south during the Bell Beaker phenomenon.

The expansion of R1b1a1a2a1a-L151 lineages could then be linked to the first introduction of Indo-European languages in Western Europe[Cassidy et al. 2016], which could have left some traces of their presence in historic times. However, the later expansion of Celtic languages, and an apparent resurgence of the probably indigenous Proto-Iberian and Proto-Basque languages – possibly the descendant of the languages of early farmers, similar to Paleo-Sardinian[Terradas et al. 2014] – over an expanding Iberian subclade (R1b1a1a2a1a2a1b1a1-M167/SRY2627) of the R1b1a1a2a1a2a-DF27 lineage[Gunther et al. 2015] have left scarce data on the older situation.

Samples from northern and south-western Iberia in the transition from the Chalcolithic to the Bronze Age have been found to have steppe ancestry, although in lesser proportion than in central and western Europe[Olalde et al. 2017][Martiniano et al. 2017]. Samples of north-east Iberia, dated ca. 2385 BC, show haplogroups R1b1a-L754 (xR1b1a1a2a-L23), G2-P287, and I2a2-M436, which point to at least a partial persistence of old European hunter-gatherer lineages in the region.

However, there is a clear replacement of autochthonous I2-M438 and G2a-P15 subclades by R1b1a1a2a1a2-P312. The expansion of haplogroup R1b1a1a2a1a2a-DF27 from northern Iberia, based on studies of the modern population, also points to ca. 2200 BC[Solé-Morata et al. 2017]. The greater admixture with the autochthonous population probably suggests a different type of expansion than that of Bell Beakers in central Europe and the British Isles, probably a later renewed male-driven invasion from northern Iberia, once admixed with the local population.

The only certain Indo-European language of Iberia that can be considered of a non-Celtic nature is Lusitanian (which has been linked to a potential Galaico-Lusitanian group of the north-western Iberian Peninsula), and there has been some discussion on the pre-Celtic nature of the languages of Cantabri, Astures, Pellendones, Carpetani, and Vettones. Also, while the position of Tartessian as Indo-European[Koch 2009] is highly doubted[2] – , there is some support for a borrowing of names from a “lost Indo-European language” over the course of long-term contacts[Mikhailova 2015].

The emergence of El Argar groups was preceded by a break in Chalcolithic cultural traditions in south-east Iberia, which points to an upheaval of existing social structures or an influx of groups that cannot be distinguished from the local population at the present of genetic resolution, e.g. from south-eastern Europe[Szecsenyi-Nagy et al. 2017]. This could point to the time of resurge of groups associated with previous Neolithic cultures that might have conserved Pre-Iberian and Pre-Basque languages until historic times.

The other region where modern R1b1a1a2a1a2a-DF27 lineages peak in the modern population corresponds to the old Nordwestblock cultural region, where a non-Celtic, non-Germanic Indo-European language might have been spoken[Kuhn, Hachmann, and Kossack 1986].

r1b-DF27.jpg Modern distribution of haplogroup R1b1a1a2a1a2a-DF27. Adapted from Myres et al.[Myres et al. 2011]

To the east, in the Vistula group of Lesser Poland, Bell Beaker samples of R1b1a1a2-M269 are found from 2400 BC to 2300 BC. Later an eastern sample is found near the Oder ca. 2170 BC, which – together with long-term and long-distance economic exchange (especially regarding amber imports) during the Bronze Age[Makarowicz 2009] – support the presence of Old European river names in east Europe dating to this period.


  • [Adrados 1998] ^ Adrados, F.R. 1998. La reconstrucción del indoeuropeo y de su diferenciación dialectal. In Manual de lingüística indoeuropea, edited by F. R. Adrados, A. Bernabé and J. Mendoza. Madrid: Ediciones clásicas.
  • [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.
  • [Brodie 2001] ^ Brodie, N. 2001. Technological frontiers and the emergence of the Beaker Culture. In Bell Beakers today: pottery, people, culture, symbols in prehistoric Europe, edited by F. Nicolis. Trento: Servicio Beni Culturali, Provincia Autonoma di Trento.
  • [Cassidy et al. 2016] ^ 1 2 Cassidy, L. M., R. Martiniano, E. M. Murphy, M. D. Teasdale, J. Mallory, B. Hartwell, and D. G. Bradley. 2016. Neolithic and Bronze Age migration to Ireland and establishment of the insular Atlantic genome. Proc Natl Acad Sci U S A 113 (2):368-73.
  • [Fokkens and Nicolis 2012] ^ Fokkens, Harry, and Franco Nicolis. 2012. Background to Beakers: Inquiries in Regional Cultural Backgrounds of the Bell Beaker Complex. Leiden: Sidestone Press.
  • [Gunther et al. 2015] ^ Gunther, T., C. Valdiosera, H. Malmstrom, I. Urena, R. Rodriguez-Varela, O. O. Sverrisdottir, E. A. Daskalaki, P. Skoglund, T. Naidoo, E. M. Svensson, J. M. Bermudez de Castro, E. Carbonell, M. Dunn, J. Stora, E. Iriarte, J. L. Arsuaga, J. M. Carretero, A. Gotherstrom, and M. Jakobsson. 2015. Ancient genomes link early farmers from Atapuerca in Spain to modern-day Basques. Proc Natl Acad Sci U S A 112 (38):11917-22.
  • [Harrison and Heyd 2007] ^ 1 2 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] ^ 1 2 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.
  • [Knipper et al. 2017] ^ Knipper, Corina, Alissa Mittnik, Ken Massy, Catharina Kociumaka, Isil Kucukkalipci, Michael Maus, Fabian Wittenborn, Stephanie E. Metz, Anja Staskiewicz, Johannes Krause, and Philipp W. Stockhammer. 2017. Female exogamy and gene pool diversification at the transition from the Final Neolithic to the Early Bronze Age in central Europe. Proceedings of the National Academy of Sciences.
  • [Koch 2009] ^ Koch, John T. 2009. A case for tartessian as a celtic language. Palaeohispanica 9:339-351.
  • [Krahe 1949] ^ Krahe, H. 1949. Alteuropäische Flußnamen. Beiträge zur Namenforschung 1:24–51, 247–266 (and cont. in following volumes).
  • [Krahe 1964] ^ Krahe, H. 1964. Unsere ältesten Flußnamen. Wiesbaden: Harrassowitz.
  • [Kuhn, Hachmann, and Kossack 1986] ^ Kuhn, Hans, Rolf Hachmann, and Georg Kossack. 1986. Völker zwischen Germanen und Kelten. Schriftquellen, Bodenfunde und Namengute zur Geschichte des nördlichen Westdeutschlands um Christi Gebur. Neumünster: Karl Wachholz.
  • [Lazaridis et al. 2016] ^ Lazaridis, I., D. Nadel, G. Rollefson, D. C. Merrett, N. Rohland, S. Mallick, D. Fernandes, M. Novak, B. Gamarra, K. Sirak, S. Connell, K. Stewardson, E. Harney, Q. Fu, G. Gonzalez-Fortes, E. R. Jones, S. A. Roodenberg, G. Lengyel, F. Bocquentin, B. Gasparian, J. M. Monge, M. Gregg, V. Eshed, A. S. Mizrahi, C. Meiklejohn, F. Gerritsen, L. Bejenaru, M. Bluher, A. Campbell, G. Cavalleri, D. Comas, P. Froguel, E. Gilbert, S. M. Kerr, P. Kovacs, J. Krause, D. McGettigan, M. Merrigan, D. A. Merriwether, S. O'Reilly, M. B. Richards, O. Semino, M. Shamoon-Pour, G. Stefanescu, M. Stumvoll, A. Tonjes, A. Torroni, J. F. Wilson, L. Yengo, N. A. Hovhannisyan, N. Patterson, R. Pinhasi, and D. Reich. 2016. Genomic insights into the origin of farming in the ancient Near East. Nature 536 (7617):419-24.
  • [Lee et al. 2012] ^ Lee, E. J., C. Makarewicz, R. Renneberg, M. Harder, B. Krause-Kyora, S. Muller, S. Ostritz, L. Fehren-Schmitz, S. Schreiber, J. Muller, N. von Wurmb-Schwark, and A. Nebel. 2012. Emerging genetic patterns of the European Neolithic: perspectives from a late Neolithic Bell Beaker burial site in Germany. Am J Phys Anthropol 148 (4):571-9.
  • [Makarowicz 2009] ^ Makarowicz, Przemysław. 2009. Baltic-Pontic Interregional Routes at the Start of the Bronze Age. Baltic-Pontic Studies 14:301-336.
  • [Mathieson et al. 2017] ^ 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. https://doi.org/10.1101/135616
  • [Mikhailova 2015] ^ Mikhailova, Tatyana A. 2015. Celtic origin: location in time and space? Reconsidering the “East-West Celtic” debate. Journal of Language Relationship 13 (3):257-279.
  • [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. https://doi.org/10.1101/113241
  • [Myres et al. 2011] ^ 1 2 Myres, N. M., S. Rootsi, A. A. Lin, M. Jarve, R. J. King, I. Kutuev, V. M. Cabrera, E. K. Khusnutdinova, A. Pshenichnov, B. Yunusbayev, O. Balanovsky, E. Balanovska, P. Rudan, M. Baldovic, R. J. Herrera, J. Chiaroni, J. Di Cristofaro, R. Villems, T. Kivisild, and P. A. Underhill. 2011. A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe. Eur J Hum Genet 19 (1):95-101.
  • [Nicolaisen 1957] ^ Nicolaisen, W. 1957. Die alteuropäischen Gewässernamen der britischen Hauptinsel. Beiträge zur Namenforschung:211-268.
  • [Olalde et al. 2017] ^ 1 2 3 4 5 6 Olalde, Iñigo, Selina Brace, Morten E. Allentoft, Ian Armit, Kristian Kristiansen, Nadin Rohland, Swapan Mallick, Thomas Booth, Anna Szécsényi-Nagy, Alissa Mittnik, Eveline Altena, Mark Lipson, Iosif Lazaridis, Nick J. Patterson, Nasreen Broomandkhoshbacht, Yoan Diekmann, Zuzana Faltyskova, Daniel M. Fernandes, Matthew Ferry, Eadaoin Harney, Peter de Knijff, Megan Michel, Jonas Oppenheimer, Kristin Stewardson, Alistair Barclay, Kurt W. Alt, Azucena Avilés Fernández, Eszter Bánffy, Maria Bernabò-Brea, David Billoin, Concepción Blasco, Clive Bonsall, Laura Bonsall, Tim Allen, Lindsey Büster, Sophie Carver, Laura Castells Navarro, Oliver Edward Craig, Gordon T. Cook, Barry Cunliffe, Anthony Denaire, Kirsten Egging Dinwiddy, Natasha Dodwell, Michal Ernée, Christopher Evans, Milan Kuchařík, Joan Francès Farré, Harry Fokkens, Chris Fowler, Michiel Gazenbeek, Rafael Garrido Pena, María Haber-Uriarte, Elżbieta Haduch, Gill Hey, Nick Jowett, Timothy Knowles, Ken Massy, Saskia Pfrengle, Philippe Lefranc, Olivier Lemercier, Arnaud Lefebvre, Joaquín Lomba Maurandi, Tona Majó, Jacqueline I. McKinley, Kathleen McSweeney, Mende Balázs Gusztáv, Alessandra Modi, Gabriella Kulcsár, Viktória Kiss, András Czene, Róbert Patay, Anna Endródi, Kitti Köhler, Tamás Hajdu, João Luís Cardoso, Corina Liesau, Michael Parker Pearson, Piotr Włodarczak, T. Douglas Price, Pilar Prieto, Pierre-Jérôme Rey, Patricia Ríos, Roberto Risch, Manuel A. Rojo Guerra, Aurore Schmitt, Joël Serralongue, Ana Maria Silva, Václav Smrčka, Luc Vergnaud, João Zilhão, David Caramelli, Thomas Higham, Volker Heyd, Alison Sheridan, Karl-Göran Sjögren, Mark G. Thomas, Philipp W. Stockhammer, Ron Pinhasi, Johannes Krause, Wolfgang Haak, Ian Barnes, Carles Lalueza-Fox, and David Reich. 2017. The Beaker Phenomenon And The Genomic Transformation Of Northwest Europe. bioRxiv. https://doi.org/10.1101/135962
  • [Parker Pearson et al. 2016] ^ Parker Pearson, Mike, Andrew Chamberlain, Mandy Jay, Mike Richards, Alison Sheridan, Neil Curtis, Jane Evans, Alex Gibson, Margaret Hutchison, Patrick Mahoney, Peter Marshall, Janet Montgomery, Stuart Needham, Sandra O'Mahoney, Maura Pellegrini, and Neil Wilkin. 2016. Beaker people in Britain: migration, mobility and diet. Antiquity 90 (351):620-637.
  • [Prieto Martínez and Salanova 2015] ^ Prieto Martínez, Maria Pilar, and Laure Salanova. 2015. Concluding remarks. The Bell Beaker Transition: The end of the neolithisation of Europe; the starting point of a new order. In The Bell BeakerTransition in Europe. Mobility and local evolution during the 3rd millenium BC, edited by M. P. P. Martínez and L. Salanova. Oxford & Philadelphia: Oxbow Books.
  • [Szecsenyi-Nagy et al. 2017] ^ Szecsenyi-Nagy, Anna, Christina Roth, Brandt Guido, Cristina Rihuete-Herrada, Cristina Tejedor-Rodriguez, Petra Held, Inigo Garcia-Martinez-de-Lagran, Hector Arcusa Magallon, Stephanie Zesch, Corina Knipper, Eszter Banffy, Susanne Friedrich, Harald Meller, Primitiva Bueno-Ramirez, Rosa Barroso Bermejo, Rodrigo de Balbin Behrmann, Ana M. Herrero-Coral, Raul Flores Fernandez, Carmen Alonso Fernandez, Javier Jimenez Echevarria, Laura Rindlisbacher, Camila Oliart, Maria-Ines Fregeiro, Ignacio Soriano, Oriol Vincente, Rafael Mico, Vincente Lull, Jorge Soler Diaz, Juan Antonio Lopez Padilla, Consuelo Roca de Togores Munoz, Mauro S. Hernandez Perez, Francisco Javier Jover Maestre, Joaquin Lomba Maurandi, Azucena Aviles Fernandez, Katina T. Lillios, Ana Maria Silva, Miguel Magalhaes Ramalho, Luiz Miguel Oosterbeek, Claudia Cunha, Anna J Waterman, Jordi Roig Buxo, Andres Martinez, Juana Ponce Martinez, Mark Hunt Ortiz, Juan Carlos Mejias-Gracia, Juan Carlos Pecero Espin, Rosario Cruz-Aunon Briones, Tiago Tome, Eduardo Carmona Ballestero, Joao Luis Cardoso, Ana Cristina Araujo, Corina Liesau von Lettow-Vorbeck, Conception Blasco Bosqued, Patricia Rios Mendoza, Ana Pujante, Jose I. Royo-Guillen, Marco Aurelio Esquembre Bevia, Victor Manuel Dos Santos Goncalves, Rui Parreira, Elena Moran Hernandez, Elena Mendez Izquierdo, Jorge Vega de Miguel, Roberto Menduina Garcia, Victoria Martinez Calvo, Oscar Lopez Jimenez, Johannes Krause, Sandra L. Pichler, Rafael Garrido-Pena, Michael Kunst, Roberto Risch, Manuel A. Rojo-Guerra, Wolfgang Haak, and Kurt W. Alt. 2017. The maternal genetic make-up of the Iberian Peninsula between the Neolithic and the Early Bronze Age. bioRxiv.
  • [Terradas et al. 2014] ^ Terradas, Xavier, Bernard Gratuze, Josep Bosch, Roser Enrich, Xavier Esteve, F. Xavier Oms, and Genís Ribé. 2014. Neolithic diffusion of obsidian in the western Mediterranean: new data from Iberia. Journal of Archaeological Science 41:69-78.
  • [Vander Linden 2007] ^ Vander Linden, Marc. 2007. For equalities are plural: reassessing the social in Europe during the third millenniumbc. World Archaeology 39 (2):177-193.


  1. ZZ11+ equivalent (ancestral to DF27 and U152); DF27+? In 390k BAM file, but short, may actually belong to chromosomes 2 or 5. Additional information from Alex Williamson.
  2. It was criticized extensively in a special section of Vol. 42 of The Journal of Indo-European Studies (No. 3 & 4, Fall/Winter 2014)