View source for Caucasus Hunter-Gatherer ancestry and Indo-Hittite

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Caucasus hunter-gatherer (CHG) ancestry is defined by a Late Palaeolithic individual from Satsurblia cave (ca. 11000 BC), and a Mesolithic individual from Kotias Klde cave, in western Georgia (ca. 6000 BC) – both of haplogroup J-M304 (the Kotias sample, and possibly both, from J2-M172). 

Caucasus hunter-gatherers seem to have weathered much of the last Ice Age in isolation, with the individual from Satsurblia showing also signs of recent consanguinity. That isolation has continued partially into the modern population of the southern Caucasus, in terms of ancestry, as well as Y-DNA and mtDNA haplogroups[Jones et al. 2015]. 

CHG ancestry was believed until recently to have contributed only late to the population of the Pontic-Caspian steppe, coinciding with the formation of the Yamna culture, since Samara hunter-gatherers showed only EHG ancestry and no CHG ancestry, while Yamna samples had up to 43% of CHG ancestry[Jones et al. 2015][Lazaridis et al. 2016]. However, steppe ancestry accompanied by CHG component has been found in an individual from the Balkans at Varna I (ca. 4630 BC), and steppe ancestry in Smyadovo (ca. 4500 BC), some 2,000-1,000 years before the main Chalcolithic expansions associated with the steppes[Mathieson et al. 2017]. 

<span class="plainlinks">[https://indo-european.eu/human-ancestry/admixture-ancestry-components-r-statistics-plink-convertf-bed-and-ped-files/ https://indo-european.eu/wp-content/uploads/2017/10/steppe-component.png]</span>

<small>''Modified from [https://doi.org/10.1101/135616 Mathieson et al (2017)]. «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 (orange), EHG (red) and WHG (blue)».  Dates indicate approximate range of individuals in each population[Mathieson et al. 2017]. Original image under a [http://creativecommons.org/licenses/by-nc/4.0/ CC-BY-NC 4.0 International license].''.</small>

<span class="plainlinks">[https://indo-european.eu/human-ancestry/principal-component-analysis-pca-and-eigensoft/ https://indo-european.eu/wp-content/uploads/2017/10/pca2-mathieson.png]</span>

<small>''Modified from [https://doi.org/10.1101/135616 Mathieson et al (2017)]. «Individuals projected onto axes defined by the principal components of 799 present-day West Eurasians (not shown in thisplot for clarity, but shown in Extended Data Figure 1). Projected points include selected published individuals (faded colored circles, labeled) and newly reported individuals (other symbols; outliers shown by additional black circles). Colored polygons indicate the individuals that had cluster memberships fixed at 100% for the supervised admixture analysis [on the right]».</small>

This suggests a more likely gradual contribution from intermittent contacts with cultures from the Caucasus during the Mesolithic-Neolithic and Neolithic-Chalcolithic transitions, joint with the westward expansions (and probably inner west-east movements) of Middle Indo-European speakers in the Pontic-Caspian steppe and beyond its natural frontiers to the north. 

This CHG ancestry might have been involved in what is believed to be a Caucasian substratum of Indo-European[Bomhard 2017], potentially differentiating it from Uralic within a hypothetic ancient Indo-Uralic group (Kortlandt 2002). On the other hand, CHG ancestry has also been found in Neolithic samples from the north Pontic steppe regions, so that the linguistic differences between Uralic and Indo-Hittite cannot be explained solely by the contribution of the population from the Caucasus.

<span class="plainlinks">[https://indo-european.eu/maps/mesolithic/ https://indo-european.eu/wp-content/uploads/2017/09/mesolithic_Steppe.jpg]</span>
''Diachronic map of Mesolithic migrations ca. 6500-5000 BC [Anthony 2007][Piezonka 2015], [http://context-database.uni-koeln.de/ Uni-Köln].''

<span class="plainlinks">[https://indo-european.eu/maps/neolithic/ https://indo-european.eu/wp-content/uploads/2017/05/neolithic_steppe.jpg]</span>
''Diachronic map of Early Neolithic migrations ca. 5000-4000 BC near the Caucasus [Anthony 2007][Piezonka 2015][Czekaj-Zastawny, Kabaciński, and Terberger 2015].''

== References ==

* [*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.

* [*Bomhard 2017] Bomhard, Alan R. 2017. [https://www.academia.edu/10261406/The_Origins_of_Proto-Indo-European_The_Caucasian_Substrate_Hypothesis_revised_August_2017_ The Origins of Proto-Indo-European: The Caucasian Substrate Hypothesis]. Charleston, SC.

* [*Czekaj-Zastawny, Kabaciński, and Terberger 2015] Czekaj-Zastawny, Agnieszka, Jacek Kabaciński, and Thomas Terberger. 2015. Long distance exchange in the Central European Neolithic: Hungary to the Baltic. Antiquity 85 (327):43-58.

* [*Jones et al. 2015] Jones, E. R., G. Gonzalez-Fortes, S. Connell, V. Siska, A. Eriksson, R. Martiniano, R. L. McLaughlin, M. Gallego Llorente, L. M. Cassidy, C. Gamba, T. Meshveliani, O. Bar-Yosef, W. Muller, A. Belfer-Cohen, Z. Matskevich, N. Jakeli, T. F. Higham, M. Currat, D. Lordkipanidze, M. Hofreiter, A. Manica, R. Pinhasi, and D. G. Bradley. 2015. Upper Palaeolithic genomes reveal deep roots of modern Eurasians. Nat Commun 6:8912. https://doi.org/10.1038/ncomms9912

* [*Kortlandt 2002] Kortlandt, Frederik. 2002. [http://www.kortlandt.nl/publications/art203e.pdf The Indo-Uralic verb]. In Finno-Ugrians and Indo-Europeans: Linguistic and literary contacts. Maastricht: Shaker.

* [*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. [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5003663/ Genomic insights into the origin of farming in the ancient Near East]. Nature 536 (7617):419-24. https://dx.doi.org/10.1038/nature19310

* [*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

* [*Piezonka 2015] Piezonka, Henny. 2015. Older than the farmers' pots? Hunter-gatherer ceramics east of the Baltic Sea. In The Dąbki Site in Pomerania and the Neolithisation of the North European Lowlands (c. 5000-3000 calBC), edited by J. Kabaciński, S. Hatz, R. D. C. M. and T. Terberger. Rahden/Westf.: Marie Leidorf.

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