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. 2019 Aug 13;9:11759. doi: 10.1038/s41598-019-47972-1

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Above (right): Schematic map of the main IUP sites in Central Europe (Bohunician) (A), in the Middle-East (Emirean) (B) and in Asia (D); and the Ust-Ishim human femur (C). Above (left): View on the T16 site and the Western flank of Tolbor Valley. Below: Detailed map of the potential IUP sites in the region. 1, Luotuoshi; 2, Ush-Bulak; 3, Malo Yaloman, Cave; 4, Kara-Bom, 5, Ust-Karakol-1 and Denisova Cave; 6, Kara-Tenesh; 7, Derbina sites; 8, Arembovski; 9, Makarovo-IV; 10, Khotyk; 11, Barun-Alan sites; 12, Kamenka and Varvarina Gora; 13, Tolbaga; 14, Podzvonkaya; 15, Egiin-Gol sites (Dörölj 1–2); 16, Tolbor-16 and Tolbor-4; 17, Tsatsyn Ereg; 18, Mojlt’yn-Am; 19, Chiken sites; 20, Tsagan-Agui; 21, Shuiddonggou 1; 22, Khanzat-1; 23, Khavsgayt (and Salkhit); 24, Rashaan Khad; 25, Otson Tsokhio^{16–18,75–77}; Geo-atlas background map).