The southern stream of Yamna migrants showed a later expansion from the Lower Danube and Tisza rivers to the southern tip of the Balkan peninsula, and its population is probably represented by the transition of Early Helladic II to Early Helladic III period ca. 2200 BC[Gimbutas 1977], coincident with the arrival of Minyan pottery style. Others have proposed a later date, the beginning of the Middle Helladic culture ca. 2000-1900 BC[Beekes 2011].
A recent study of Minoan samples from Crete (ca. 2900-1700 BC) and Mycenaean samples from mainland southern Greece (ca. 1700-1200 BC) have shown an introduction of CHG without EHG compared to Mesolithic samples, thus independent of a steppe invasion. This eastern influence is also found in Y-DNA haplogroups, all four of them J-M304, related to Bronze Age pastoralists of the Eurasian steppe. This eastern influence may have arrived from Anatolia, since Neolithic samples from central Anatolia at Tepecik-Çiftlik already show it some millennia earlier[Kilinc et al. 2016], and more recent samples from south-western Anatolian (ca. 2800-1800 BC) show it too.
Mycenaean samples show a ‘northern’ contribution, apart from the Neolithic farmer and ‘eastern’ ancestry shown by Minoan samples. This ‘eastern’ contribution may be interpreted in part as a steppe ancestry similar to that found in the Balkans in the third millennium, suggesting a rapid migration of Proto-Greek from the Balkans, although it is found only in samples from mainland Greece, and not in one sample from Crete[Lazaridis et al. 2017].
The introduction of millet and the horse in northern Greece seem to coincide, pointing to their introduction by horse breeding, millet-consuming cultures from the north or north-east, via river valleys leading to the Danube[Valamoti 2016].
The potential invasion and assimilation of R1b1a1a2a2-Z2103 settlers in Proto-Anatolian-speaking populations in Cernavodă III and Ezero cultures further confounds modern genetic studies, and more aDNA samples are needed to more clearly depict the expansion of both populations. Such a contact may have happened early during the southward migration, as suggested by Anatolian loanwords found in Greek.
The complexity of Y-DNA haplogroups found in the modern population of Greece bears witness to the thousands of years of European and Asian interaction in the formation of its peoples. While it is clear that Yamna ancestry does not represent a big part of its modern population in any study published to date, it is unclear how the ancient population was affected by the migration of peoples of R1b1a1a2a2-Z2103 lineages.
[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 Potapovka cultures, as well as Bell Beaker.PCA analysis of free datasets including Minoans and Mycenaeans
Analysis of modern Greek and Cretan lineages point to a Neolithic expansion of haplogroup R1b1a1a2-M269 in the region, which was found nearer to Italian than to Balkan lineages[King et al. 2008]. Analysis of Greek-Cypriot modern populations revealed the presence of R1b1a1a2a2-Z2103 lineages in easternmost and westernmost sides of the island, with central (R1b1a1a2a2c1a-Z2110) lineages appearing only in the east[Voskarides et al. 2016]. The early attestation of Mycenaean Greek in the island point to an early expansion of R1b1a1a2a2c1a-Z2110 lineages, but the early Anatolian influence over the island precludes a precise identification of their origin.
Most R1b1a1a2-M269 lineages in Greece are probably linked to the Bronze Age expansion associated with the Minyan pottery, in turn linked to an earlier Yamna expansion into south-eastern Europe. Most R1a1a1b2-Z93 lineages – whose proportion in Greece and in historically Greek Anatolia increases with latitude and in peripheral areas – are thus to be linked to the recent southern migration of Balkanic populations of R1a1a1b1-Z283 lineages (mainly Slavs, but also Aromanians, and Albanians), and to the western spread of Iranian peoples of R1a1a1b2-Z93 lineages[Heraclides et al. 2017].
A potentially older invasion of certain R1a1a1b2-Z93 lineages during the Bronze Age could be supported by the finding of haplogroup R1a1a1b2-Z93 in an individual from Merichleri, in Bronze Age Bulgaria, ca. 1690 BC[Mathieson et al. 2017].
On the controversial ethnicity and language of the Sea Peoples and the closely related Philistine question [Woudhuizen 2006][Maeir, Davis, and Hitchcock 2016][Middleton 2015], genetic research points to a mixture of steppe ancestry found in the Lebanese population that occurred ca. 1740-160 BC [Haber et al. 2017] – probably in the earlier part of that period, possibly still earlier than the estimated range –, showing a clear haplogroup turnover in the modern Levantine population, with new subclades R1b1a1a2a2-Z2103 potentially from early Balkan (IE) individuals, and R1a1a1b2-Z93 probably from eastern (Iranian) migrants.
- [Anthony 2007] ^ 1 2 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.
- [Beekes 2011] ^ Beekes, Robert S.P. 2011. Comparative Indo-European Linguistics. An introduction. 2nd ed. Amsterdam / Philadelphia: John Benjamins.
- [Butler, Arnoldussen, and Steegstra 2011/2012] ^ Butler, J.J., S. Arnoldussen, and H. Steegstra. 2011/2012. Single-edged socketed Urnfield knives in the Netherlands and western Europe. Palaeohistoria 53/54:65-107.
- [Fokkens and Harding 2013] ^ 1 2 Fokkens, Harry, and Anthony Harding. 2013. The Oxford Handbook of European Bronze Age. Oxford: Oxford University Press.
- [Gimbutas 1977] ^ Gimbutas, Marija. 1977. The first wave of eurasian pastoralists into copper age europe. JIES 5 (4):277-338.
- [Haber et al. 2017] ^ Haber, Marc, Claude Doumet-Serhal, Christiana Scheib, Yali Xue, Petr Danecek, Massimo Mezzavilla, Sonia Youhanna, Rui Martiniano, Javier Prado-Martinez, Michał Szpak, Elizabeth Matisoo-Smith, Holger Schutkowski, Richard Mikulski, Pierre Zalloua, Toomas Kivisild, and Chris Tyler-Smith. 2017. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. bioRxiv.
- [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.
- [Heraclides et al. 2017] ^ Heraclides, Alexandros, Evy Bashiardes, Eva Fernández-Domínguez, Stefania Bertoncini, Marios Chimonas, Vasilis Christofi, Jonathan King, Bruce Budowle, Panayiotis Manoli, and Marios A. Cariolou. 2017. Y-chromosomal analysis of Greek Cypriots reveals a primarily common pre-Ottoman paternal ancestry with Turkish Cypriots. PLOS ONE 12 (6):e0179474.
- [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.
- [King et al. 2008] ^ King, R. J., S. S. Ozcan, T. Carter, E. Kalfoglu, S. Atasoy, C. Triantaphyllidis, A. Kouvatsi, A. A. Lin, C. E. Chow, L. A. Zhivotovsky, M. Michalodimitrakis, and P. A. Underhill. 2008. Differential Y-chromosome Anatolian influences on the Greek and Cretan Neolithic. Ann Hum Genet 72 (Pt 2):205-14.
- [Kilinc et al. 2016] ^ Kilinc, G. M., A. Omrak, F. Ozer, T. Gunther, A. M. Buyukkarakaya, E. Bicakci, D. Baird, H. M. Donertas, A. Ghalichi, R. Yaka, D. Koptekin, S. C. Acan, P. Parvizi, M. Krzewinska, E. A. Daskalaki, E. Yuncu, N. D. Dagtas, A. Fairbairn, J. Pearson, G. Mustafaoglu, Y. S. Erdal, Y. G. Cakan, I. Togan, M. Somel, J. Stora, M. Jakobsson, and A. Gotherstrom. 2016. The Demographic Development of the First Farmers in Anatolia. Curr Biol 26 (19):2659-2666.
- [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 2000] ^ Kristiansen, K. 2000. Europe Before History. Cambridge: Cambridge University Press.
- [Kristiansen 2014] ^ Kristiansen, K. 2014. Bronze Age Identities. In A Companion to Ethnicity in the Ancient Mediterranean, edited by J. McInerney: Wiley-Blackwell.
- [Kristiansen 2016] ^ Kristiansen, K. 2016. Bronze Age Vikings? A Comparative Analysis of Deep Historical Structures and their Dynamics. In Comparative Perspectives on Past Colonisation, Maritime Interaction and Cultural Integration. New Directions in Anthropological Archaeology, edited by L. Melheim, Z. T. Glørstad and H. Glørstad. Sheffield: Equinox.
- [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.
- [Lazaridis et al. 2017] ^ 1 2 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.
- [Maeir, Davis, and Hitchcock 2016] ^ Maeir, Aren M., Brent Davis, and Louise A. Hitchcock. 2016. Philistine Names and Terms Once Again: A Recent Perspective. Journal of Eastern Mediterranean Archaeology & Heritage Studies 4 (4):321-340.
- [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.
- [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).
- [Middleton 2015] ^ Middleton, Guy D. 2015. Telling Stories: The Mycenaean Origins of the Philistines. Oxford Journal of Archaeology 34 (1):45-65.
- [Przybyła 2009] ^ 1 2 Przybyła, Marcin S. 2009. Intercultural Contacts in the Western Carpathian Area at the Turn of the 2nd and 1st Millennia BC: Narodowe Centrum Kultury.
- [Przybyła 2016] Valamoti, Soultana Maria. 2016. Millet, the late comer: on the tracks of Panicum miliaceum in prehistoric Greece. Archaeological and Anthropological Sciences 8 (1):51-63.
- [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. https://doi.org/10.1038/ncomms14615
- [Voskarides et al. 2016] ^ Voskarides, K., S. Mazieres, D. Hadjipanagi, J. Di Cristofaro, A. Ignatiou, C. Stefanou, R. J. King, P. A. Underhill, J. Chiaroni, and C. Deltas. 2016. Y-chromosome phylogeographic analysis of the Greek-Cypriot population reveals elements consistent with Neolithic and Bronze Age settlements. Investig Genet 7:1.
- [Wels-Weyrauch 2011] ^ 1 2 Wels-Weyrauch, Ulrike. 2011. Colliers nur zu Zierde? In Bronzen im Spannungsfeld zwischen praktischer Nutzung under symbolischer Bedeutung: Praehistorische Bronzefunde, Abtailung XX, 13 Band, edited by U. L. Dietz and A. Jockenhövel. Stuttgart: Franz Steiner Verlag.
- [Woudhuizen 2006] ^ Woudhuizen, Frederik Christiaan. 2006. The ethnicity of the Sea Peoples, College voor Promoties, Erasmus Universiteit Rotterdam, Rotterdam.