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1 Supplementary Material for:

2 The origin of the : genomic evidence from a 36,000--old Eastern

3 European

4 E. Andrew Bennett, Sandrine Prat, Stéphane Péan, Laurent Crépin, Alexandr Yanevich, Simon

5 Puaud, Thierry Grange, Eva-Maria Geigl

6

7

8 Table of Contents

9 Supplementary Text

10 1. Cultural context of the technocomplex 2

11 2. Buran Kaya III site, chronological, cultural and anthropological information 5

12 3. Strategy of collection and sampling protocol 12

13 Supplementary Text References 15

14 Supplementary Tables

15 Table S1. Comparative 14C dates 21

16 Table S2. Shotgun library results 22

17 Table S3. Y SNP summary with Palaeolithic comparisons 23

18 Table S4. content comparison 24

19 Table S5. Mitochondrial sequence references 25

20 Table S6. Genome sequence references 27

21 Table S7. f3(BuranKaya3A, x, Mbuti) results 29

22 Supplementary Tables References 31

23

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24 Supplementary Text 1. Cultural context of the Gravettian technocomplex

25 By Stephane Péan

26 UMR 7194 (HNHP), MNHN/CNRS/UPVD, Muséum national d’Histoire naturelle, Alliance

27 Sorbonne Université, Institut de Paléontologie Humaine, Paris,

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29 Definition and general chronology

30 The Gravettian is defined as a mid-Upper Palaeolithic, European-wide, technocomplex of cultural

31 traditions1–3, named after the site La Gravette (France), where specific lithic straight-backed points

32 (pointe de la Gravette) were first described4.

33 The Gravettian lithic can be characterized by elaborated, light, regular and precise

34 débitage and straight-backed pointed , which probably represent projectile

35 implements2,5. The Gravettian cultural facies present also characteristic osseous industry artefacts,

36 personal ornaments, portable (e.g. female figurines), or spatially organized archaeological

37 structures made from stones or interpreted as dwelling structures at some sites.

38 The Gravettian cultures in Central have been divided into eight regional facies2 and three

39 main chronological phases6–9: early phase, characterised by predominant leaf-shaped points and

40 fléchettes, in the Middle Danube valley (Austria, in Eastern ); middle

41 phase, characterised by abundant Gravette points and microgravettes and the appearance of new

42 microlithic and para-geometric implements, in Lower Austria, Moravia, and northeastern

43 Hungary, culturally related to Eastern European sites in the Middle Dniester and Don valleys; and

44 late phase, characterised by shouldered points, in the middle Danube valley (Lower Austria,

45 Moravia, western Slovakia), Southern Poland, middle Dniester valley, Upper Dnieper valley and

46 the Middle Don and Seym valleys, in eastern Europe.

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47 The Gravettian technocomplex chronologically spans from 32 to 21 ka 14C BP(ca. 36-23 ka cal

48 BP)10,11 (see below for early Gravettian sites). During late MIS 2, Gravettian evolves towards

49 facies, which last until 13.5 ka 14C BP (16.4-16.1 ka cal BP).

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51 Origin of the Gravettian cultural facies

52 Central Europe includes Gravettian settlements dated as early as ca. 32-30 ka 14C BP: in South

53 Poland, at 32,400 ± 650 14C BP (i.e.38.3-35.1 ka cal BP) and 31,000 ± 550 14C BP (36.1-34.0 ka

54 cal BP) in Obłazowa layer VIII12, at 31,550 ± 350 14C BP (36.1-34.7 ka cal BP) in Henryków

55 15 site layer 913; in Austria, at 30,500 + 900/-800 14C BP (36.7-33 ka cal BP) in Willendorf II layer

56 514; and in , at 29,650 ± 1,320 14C BP (37.1-31.1 ka cal BP) in Molodova V layer IX15.

57 In terms of origin, different hypotheses have been proposed. Considering an internal evolution

58 within Europe, Kozlowski (2015) argues a polycentric rather than a monocentric origin to be more

59 conceivable, i.e., in the middle Danube basin (as a prominent center), as as the Dniester and

60 Prut basins, and Crimea16. In this hypothesis, the Gravettian could have evolved locally from

61 regional post- transition industries with leaf-shaped points in Eastern Europe17, such as

62 the into the Pavlovian in the Middle Danube basin18, or similar local transitional

63 industries into the Molodovian in the Dniester valley19. The Gravettian could have also evolved

64 from the in the Upper Danube valley20, the Proto-Aurignacian in the , the

65 in the Mediterranean areas or the Zwierzyniecian in the areas north and east of the

66 Carpathians19. In contrast, one hypothesis of an initial origin of the Gravettian in Central Europe

67 postulates that it may have resulted from acculturation between migrating Anatomically Modern

68 importing Aurignacian cultures and local producing leaf point industries5.

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69 An external origin of the Gravettian has also been proposed linking it to cultural facies of the Near

70 East and Northeastern , such as the , Lagaman and Dabban21.

71 The industry from Buran-Kaya III layers 6-2, 6-1 and 5-2, attributed to the Gravettian

72 technocomplex22,23, has been considered, together with the material from layers 6-5 to 6-3, as

73 having a number of characteristics similar to Early Upper Palaeolithic assemblages from the

74 northwestern Caucasus (, levels 1C, 1B and 1A) and southern Caucasus

75 (Dzudzuana cave , layer D; Ortvale Klde rockshelter, layers 4d and 4c)24.

76 The last 14C analyses of the three layers 6-2, 6-1 and 5-2 from Buran-Kaya III, radiometrically

77 dated between 32.5+250-230 and 29.4+190-180 ka 14C BP (37.1-33.2 ka cal BP)25,26, confirm the

78 hypothesis of a very early occurrence of Gravettian settlements prior to 30 ka BP in Eastern

79 Europe16,27.

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92 Supplementary Text 2. Buran Kaya III site, chronological, cultural and anthropological

93 information

94 By Sandrine Prat

95 UMR 7194 (HNHP), MNHN/CNRS/UPVD, Alliance Sorbonne Université, Musée de l’Homme,

96 Paris, France

97

98 Chronological, cultural context and settlement patterns

99 Buran-Kaya III is a located on the eastern part of Crimea, in the Belogorsk region, in

100 the middle basin of the Burulcha River (4 km south from the city of Aromatne). This site was

101 discovered in 1990 by A. Yanevich and excavated initially until 2001 by a team directed by A.

102 Yanevich ( to Palaeolithic layers) and A. Marks (Middle and Early Upper Palaeolithic

103 layers) with the participation of V. Chabai, Y. Demidenko, K. Monigal, M. Otte and Y. Yamada,

104 followed by new fieldwork conducted by A. Yanevich and S. Péan between 2009 and 2011. This

105 rockshelter unearthed an exceptional stratigraphic sequence from the Middle Palaeolithic to the

106 Neolithic which lead to multiple studies22,23,25,26,28–47.

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108 Buran-Kaya III is a key site for understanding the Middle to Upper Palaeolithic transition and the

109 arrival and dispersal of Anatomical Modern Humans (AMHs) in Europe as well their potential

110 biological and/or cultural interactions with Neanderthals. During the excavation season of 2001

111 and 2009-2011, this site has yielded a Middle Palaeolithic layer (Micoquian, Kiik-Koba type, layer

112 B) located above an Early Upper layer (Streletskian or eastern Szeletian, layer C) and

113 below three Aurignacian layers (layers 6-5 to 6-3), followed by three Gravettian layers (layers 6-

114 2, 6-1 and 5-2) and one Swiderian layer (layer 4), all densely layered across 1 to 2 meters of depth.

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115 The stratigraphical and radiochronological analyses26 had emphasized an early chronological

116 framework for the Middle to period in southeastern Europe. These studies show

117 the presence of Early Upper Paleolithic (early Streletskian or eastern Szeletian, layer C)

118 populations in Crimea, before 40.0 ka cal BP confirming a previous dating of 44.3–38.5 ka cal BP

119 (36.7 ± 1500 ka BP, OxA-6868) of the Streletskaya or eastern Szeletian layer30 and providing early

120 evidence of this transitional industry. Based on the new radiocarbon dates, the late Middle

121 Palaeolithic settlements (Micoquian layer, layer B) range between 35,390 +290/-270 and 37,700

122 ± 900 BP26 (43,545 and 39,300 cal BP), showing that no Neanderthal settlements occurred at

123 Buran-Kaya III after the Campanian Ignimbrite eruption around 39 ka cal BP. The two Upper

124 Paleolithic cultural traditions, related to the Aurignacian and Gravettian complexes (layers 6-5 to

125 5-2), were successively present in Crimea during a relatively short time span between 39.4 and

126 34.1 ka cal BP. During the following period of about 20,000 , no depositional processes

127 occurred or were preserved and the site was presumed abandoned. The Final Paleolithic

128 populations (Swiderian, layer 4) settled in Crimea between 11.8 and 11.3 ka cal BP, i.e. during

129 MIS 1 (Fig. S1).

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131 The layers (6-2, 6-1 and 5-2) yielded a large collection of remains, and more than 28,000

132 lithic remains were discovered during the 2001 field season, representing a large spectrum of

133 types (e.g. burins, end-scrapers, and backed ). The lithic industries exhibited a high

134 percentage of microliths (more than 80%) and some microgravettes23, which are consistent with

135 the Gravettian technocomplex. They do not present Dufour or pseudo-Dufour blades, often

136 associated with Aurignacian technocomplex, which are present in layer 6-3, 6-4 and 6-5,

137 stratigraphically located below23. As shown by the sedimentological and stratigraphic analyses

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138 conducted by S. Puaud during the 2009 and 2010 field seasons, there is continuity in the

139 stratigraphic sequence from the lower layers (Aurignacian technocomplex: 6-5, 6-4, 6-3) to the

140 upper layers (Gravettian technocomplex: 6-2, 6-1 and 5-2). The attribution of the upper layers to

141 the Gravettian tradition is free from material sedimentary overlap.

142 Concerning the industry, more than 60 bone tools have been discovered in the Gravettian

143 layers corresponding to projectile points, awls, , and assegai points23. The bone

144 industry was probably imported as indicated by the absence of manufacturing waste. This

145 evidence, combined with the zooarchaeological results44, suggests a pattern of recurrent short-term

146 occupations, such as seasonal or temporary butchery camps, associated with exploitation

147 of small or medium-sized , such as saiga antelope (the main game exploited in these

148 layers). The settlement of AMHs associated with the Gravettian layers of Buran-Kaya III occurred

149 under interstadial climatic conditions, which became cooler and drier leading to an open steppe

150 environment. The open and steppe-like periglacial environment, with a local transition zone

151 between steppes and mountains located on the migration path of several species, provided a

152 diversified faunal spectrum (e.g., saiga antelope, hare, red and polar foxes, , woolly

153 , marmot, wild cat and brown bear)26,44,47. The micromammal and palynological

154 analyses36,48 reveal a steppe environment with a climatic evolution towards an increasing aridity

155 as recorded in the mammalian sequence.

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157 Human remains

158 At Buran-Kaya III, more than 160 human remains were discovered in three well-documented

159 Gravettian layers (6-2, 6-1 and 5-2). Two of them unearthed from layer 6-1 and 6-2 have been

160 directly dated to 31,900+240/-220 BP (36.3-35.2 ka cal BP), layer 6-1 (GrA-37938)25, and 32,450

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161 +250/-230 BP (37.1-35.7 ka cal BP), layer 6-2 (GrA-50457)26. They are among the oldest direct

162 evidence of AMHs in Europe in a well-documented archaeological context (Gravettian sensu lato)

163 25,26. The Gravettian specimens from Buran-Kaya III represent, together with Kostenki, Russia

164 (Kostenki 14: 33,250 +/-500 BP; 38.7-36.3 ka cal BP49 and Peştera cu Oase, (Oase 1:

165 34,290 +970/-870 BP: 36.47-41.07 ka cal BP, GrA-22810)50, the earliest occurrence of AMHs in

166 Eastern Europe. AMHs occurred later in , e.g., at Goyet Cave in (Goyet

167 Q116-1: 30,880 +170-160 BP; 35.2-34.4 ka cal BP, GrA-46175)51, in Russia, e.g., at Sunghir

168 (Sunghir SI: 28,890 +/- 430 BP; 33.9-31.8 ka cal BP, OxA-X-2464-12; Sunghir SII: 30,100+/-550;

169 35.3-33.2 ka cal BP, OxA-A-2395-6; Sunghir SIII: 30,000 +/-550 BP; 35.2-33.3 ka cal BP, OxA-

170 X-2395-7; Sunghir SIV: 29820 +/- 280 BP; 34.5-33.5, OxA-X-2462-52)49,52 and in the Czech

171 republic e.g., at Dolní Věstonice (Dolní Věstonice 16, associated with Gravettian tradition: 25,740

172 +/-210 BP (on ); 30.6-29.4 ka cal BP, GrN-15277)49.

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174 Layer 6-1 has yielded the richest assemblage of human specimens with 150 highly fragmented

175 remains (mostly cranial parts and teeth (95%) and several hand phalanges), corresponding to at

176 least five individuals45. Due to the fragmentation of the remains, the taxonomic assignment as

177 AMH could only be performed on an occipital bone fragment and the permanent teeth. Based on

178 the combinations of morphological features on these anatomical elements (Figure 2) (namely,

179 absence of an occipital “bun” or a bilaterally transverse torus; lack of a well-developed metaconid

180 and a transverse crest on the lower premolars; lack of shovelling, labial convexity and the presence

181 of well-developed lingual tubercles on the upper first incisors; the lack of a well-developed mid-

182 trigonid crest and a large anterior fovea on the lower molars), these individuals from layer 6-1 are

183 allocated to AMHs25,47.

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184 Isotopic analyses of bone collagen revealed that the diet of these individuals consisted mainly of

185 terrestrial resources, meat playing a preponderant role in the protein intake, and of plant

186 consumption46.

187 Among the human remains from the layer 6-1 (2001, 2009 and 2010 field seasons), only a few

188 bones exhibit human modifications, such as cut marks. The human skulls were intentionally

189 selected in association with post-mortem treatments of the corpses (ritual cannibalism or a specific

190 mortuary practice, such as post-mortem disarticulation processing of corpses for secondary

191 disposal)25,44,47. Based on these taphonomic observations, the specimens from layer 6-1 represent

192 the oldest Upper Palaeolithic AMHs from Eastern Europe showing post-mortem treatment of the

193 corpses.

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195 Fig. S1. Stratigraphy and chronological frameworks of Buran Kaya III site (graphic S. Puaud),

196 modified from Péan et al., 201326.

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199 Fig. S2. Remains from Buran-Kaya III (layer 6-1). From left to right, a) occipital bone; b) right

200 lower second molar; c) right first upper incisor. Scale bar = 1 cm; photos © L.Crépin/S. Prat.

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215 3. Strategy of collection and sampling protocol

216 By Laurent Crépin

217 UMR 7194 (HNHP), MNHN/CNRS/UPVD, Muséum national d’Histoire naturelle, Alliance

218 Sorbonne Université, Institut de Paléontologie Humaine, Paris, France

219

220 The excavations performed between 2009 and 2011 incorporated a multi-sampling strategy to

221 better contextualize the site of Buran-Kaya III. Sampling for sedimentological, biogeochemical,

222 and radiometric studies were performed on all Gravettian layers, and in 2009, the sampling for the

223 palaeogenomic analyses was carried out focused on the two layers that contained most of the

224 human remains (6-1 and 6-2). To this end, a one half-square meter section was excavated

225 incorporating precautions against contamination: a maximum of two excavators dug the square

226 wearing masks and gloves, and when potential human remains were discovered, one excavator put

227 on a new full body protection suit, including masks and gloves, to perform the removal of the

228 material using new tools (Fig. S3a).

229

230 This protocol was maintained over eight days, during which time layers 6-1 and 6-2 were entirely

231 cleared. In total, six bones were identified and inventoried using the described procedure, of which

232 four were certified as human remains after verification: three from layer 6-1 (two cranial fragments

233 and one tooth) and one for layer 6-2 (a cranial fragment). These remains, separated from other

234 archaeological remains, were transferred directly after the mission to the high containment

235 laboratory at the “Institut Jacques Monod” (IJM), where they were analyzed and stored at -20°C.

236 Additionally, all those working on the site during the excavation campaign provided a cheek-swab

237 sample to control for possible contamination of the samples.

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239 The anthropological and taphonomic study of the human remains was then carried out directly in

240 the high containment palaeogenomic laboratory at the IJM using typical precaution measures for

241 ancient DNA research (Fig. S3b). Finally, we decided to carry out the paleogenomics analysis on

242 the bone fragment that we could identified anatomically in an unambiguous manner that appeared

243 to be the best preserved with a minimum of visible alteration: artefact number N.135, square 9Z,

244 layer 6-1.a also called here BuranKaya3A (Fig. S3c). This piece is a posterior parietal bone

245 fragment presumably belonging to a mature individual, measuring a maximum of 21 x 19 mm.

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246

247 Fig. S3. a) Sampling of human remains during the 2009 excavation season at Buran-Kaya III;

248 photo © A. Yanevich. b) Anthropological study of human remains in the high containment

249 laboratory at the IJM; photo © E.M. Geigl. c) Images of the human parietal fragment from 6-1.a,

250 BuranKaya3A, analyzed in the present study, taken under a stereomicroscope (Leica MZ FLIII,

251 PLANAPO 0.63X); photo © E.M. Geigl.

252

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253 References

254

255 1. Garrod, D. A. E. The Upper Palaeolithic in the light of recent discovery. in 4, 1–26 (Cambridge

256 University Press, 1938).

257 2. Otte, M. Le Gravettien en Europe centrale. XX, (De Tempel, 1981).

258 3. Kozlowski, J. K. The Gravettian in Central and Eastern Europe. in Advances in world

259 archaeology 5, 131–200 (1986).

260 4. Lacorre, F. Les armatures de flèches de la Gravette: niveau et industries auxquelles elles se

261 rattachent. Procès-Verbaux Société Linéenne Bordx. 78–87 (1933).

262 5. Otte, M. Préhistoire générale. in La Préhistoire (ed. Otte, M.) 11–210 (De Boeck & Larcier,

263 1999).

264 6. Kozlowski, J. K. L’art de la préhistoire en Europe orientale. (CNRS, 1992).

265 7. Svoboda, J. & Klíma, B. Pavlov I: excavations 1952-1953. (Archeologický ústav AV ČR,

266 1994).

267 8. Otte, M., Noiret, P., Chirica, V. & Borziac, I. Rythme évolutiof du Gravettien oriental. in 213–

268 226 (ABACO, 1996).

269 9. Svoboda, J., Ložek, V. & Vlček, E. Hunters between East and West: the Paleolithic of

270 Moravia. (Springer, 1996).

271 10. Haesaerts, P., Borziac, I., Chirica, V., Damblon, F. & Koulakovska, L. Cadre stratigraphique

272 et chronologique du Gravettien en Europe centrale. PALEO Rev. Archéologie Préhistorique

273 31–51 (2007).

274 11. Noiret, P. De quoi Gravettien est-il le nom ? in Les Gravettians 29–64 (Paris: Editions Errance,

275 2013).

15

276 12. Valde-Nowak, P., Nadachowski, A., Madeyska, T. & Instytut Archeologii i Etnologii (Polska

277 Akademia Nauk). Obłazowa Cave: human activity, stratigraphy and palaeoenvironment.

278 (Institute of Archaeology and Ethnology, Polish Academy of Sciences, 2003).

279 13. Wiśniewski, A. et al. The early Gravettian in a marginal area: New evidence from SW Poland.

280 Quat. Int. 359, 131–152 (2015).

281 14. Haesaerts, P., Damblon, F., Bachner, M. & Trnka, G. Revised stratigraphy and chronology of

282 the Willendorf II sequence, Lower Austria. Archaeol. Austriaca 80, 25–42 (1996).

283 15. Sinitsyn, A., Praslov, N., Svezhentsev, Y. S. & Sulerzhitskii, L. Radiouglerodnaya

284 khronologiya verkhnego paleolita Vostochnoi Evropy. Radiouglerodnaya Khronologiya

285 Paleolita Vostochnoi Evr. Sev. Azii Probl. Perspekt. 21–66 (1997).

286 16. Otte, M. & Noiret, P. Evolution du Gravettien au moyen Danube. (Archeologický ústav AV

287 CR, 2004).

288 17. Otte, M., López Bayón, I., Noiret, P., Borziac, I. & Chirica, V. Recherches sur le Paléolithique

289 supérieur de la Moldavie. Anthropol. Praehist. Bull. Société R. Belge Anthropol. Préhistoire

290 107, 45–80 (1996).

291 18. Valoch, K. L’origine du Gravettien de l’Europe centrale. in The Upper Palaeolitic (eds. Palma

292 Di Cesnola, A., Montet-White, A. & Valoch, K.) 6, (1996).

293 19. Kozlowski, J. K. L’origine du Gravettian. in Les Gravettiens (ed. Otte, M.) 11–29 (Editions

294 Errance, 2013).

295 20. Conard, N. J. & Moreau, L. Current Research on the Gravettian of the .

296 Mitteilungen Ges. Für Urgesch. 13, 29–59 (2004).

297 21. Svoboda, J. A. The Gravettian on the Middle Danube. PALEO Rev. Archéologie Préhistorique

298 19, 203–220 (2007).

16

299 22. Yanevich, A. Buran-Kaya culture of the Crimea’s gravett. Archeologia 2, 11–20 (2000).

300 23. Yanevich, A. Les occupations gravettiennes de Buran-Kaya III (Crimée) : contexte

301 archéologique. L’Anthropologie 118, 554–566 (2014).

302 24. Demidenko, Y. E. Crimean Upper Paleolithic. in Encyclopedia of Global Archaeology (ed.

303 Smith, C.) 1782–1791 (Springer New York, 2014). doi:10.1007/978-1-4419-0465-2_1864

304 25. Prat, S. et al. The Oldest Anatomically Modern Humans from Far Southeast Europe: Direct

305 Dating, Culture and Behavior. PLoS ONE 6, e20834 (2011).

306 26. Péan, S., Puaud, S., Crépin, L., Prat, S. & Quiles, A. The Middle to Upper Paleolithic Sequence

307 of Buran-Kaya III (Crimea, Ukraine): New Stratigraphic, Paleoenvironmental, and

308 Chronological Results. Radiocarbon 55, 1454–1469 (2013).

309 27. Nuzhnyi, D. Y. The industrial variability of the eastern Gravettian assemblages of Ukraine.

310 Quartär 56, 159–174 (2009).

311 28. Yanevichv, A., Stepanchuk, V. & Cohen, Y. Buran-Kaya III and Skalistiy rockshelter: two

312 new dated Late sites in the Crimea. Préhistoire Eur. 9, 315–324 (1996).

313 29. Janevic, A. Buran-Kaya 3–Neue Angaben zur Kulturgliederung des Jungpaläolithikums der

314 Krim. Préhistoire Eur. 13, 133–148 (1998).

315 30. Marks, A. A new Middle to Upper Paleolithic “Transitional” assemblage from Buran-Kaya

316 III, level C: A preliminary report. Préhistoire Anatolie 1, 353–366 (1998).

317 31. Pettitt, P. & Otte, M. Middle and early upper Palaeolithic Crimea: the radiocarbon chronology.

318 Préhistoire D’Anatolie Genese Deux MondesAnatolian Crossroads Two Worlds 1,

319 329–338 (1998).

17

320 32. Marks, A. & Monigal, K. The Middle to Upper Palaeolithic interface at Buran-Kaya III,

321 Eastern Crimea. Neanderthals Mod. Hum.-Discuss. Transit. Cent. East. Eur. From 50–000

322 (2000).

323 33. Chabai, V. P. The Late Middle and Early Upper Paleolithic in Crimea (Ukraine). in 17, 25–36

324 (2001).

325 34. Chabai, V. The chronological and industrial variability of the Middle to Upper Paleolithic

326 transition in Eastern Europe. in 6, 71–288 (2003).

327 35. Chabai, V., Monigal, K. & Marks, A. The and early Upper Paleolithic of

328 Eastern Crimea, Volume 3 (ERAUL 104). Liège Univ. Liège (2004).

329 36. Gerasimenko, N. Vegetational of Buran-Kaya III. Paleolit. Crimea Middle Paleolit.

330 Early Up. Paleolit. East. Crimea 3, 19–34 (2004).

331 37. Markova, A. K. Small fauna from Buran-Kaya III. Paleolit. Crimea Middle Paleolit.

332 Early Up. Paleolit. East. Crimea 3, 35–48 (2004).

333 38. Monigal, K. Introduction to the site of Buran-Kaya III. Paleolit. Crimea Middle Paleolit. Early

334 Up. Paleolit. East. Crimea 3, 3–18 (2004).

335 39. Patou-Mathis, M. Archeozoological analysis of large mammal fauna from Buran-Kaya III

336 layer B. Middle Paleolit. Early Up. Paleolit. East. Crimea Liege ERAUL 104, 95–111 (2004).

337 40. Demidenko, Y. E. The Early and Mid-Upper Palaeolithic of the North region: an

338 overview. Quartär 55, 99–114 (2008).

339 41. Patou-Mathis, M. The subsistence behaviours of the Last Crimean Neanderthals. in

340 Sourcebook of Paleolithic Transitions 441–454 (Springer, 2009).

18

341 42. Yanevich, A. et al. Upper Palaeolithic settlements in Buran-Kaya 3 (Crimea, Ukraine): new

342 interdisciplinary researches of the layers 5-2, 6-1 and 6-2. in Archeological Almanac 20, 187–

343 202 (Donetsk: Donbass, 2009).

344 43. Chabai, V. The volcanic winter, Buran-Kaya cannibals and the fate of the last eastern European

345 Neanderthals. Arkheologiya 3, 5–26 (2012).

346 44. Crépin, L., Péan, S. & Lázničková-Galetová, M. Comportements de subsistance au

347 Paléolithique supérieur en Crimée: analyse archéozoologique des couches 6-2, 6-1 et 5-2 de

348 Buran-Kaya III. L’Anthropologie 118, 584–598 (2014).

349 45. Prat, S. Stress physiologique et état de santé des plus anciens Hommes anatomiquement

350 modernes du sud-est de l’Europe (données dentaires, couche 6-1, Buran-Kaya III, Crimée).

351 L’Anthropologie 118, 567–583 (2014).

352 46. Drucker, D. G. et al. Isotopic analyses suggest mammoth and plant in the diet of the oldest

353 anatomically modern humans from far southeast Europe. Sci. Rep. 7, (2017).

354 47. Prat, S. et al. The First Anatomically Modern Humans from South-Eastern Europe.

355 Contributions from the Buran-Kaya III Site (Crimea). Bull. Mém. Société Anthropol. Paris

356 (2018). doi:10.3166/bmsap-2018-0032

357 48. Gerasimenko, N. Environmental changes in the Crimean mountains during the Last

358 Interglacial–middle pleniglacial as recorded by pollen and lithopedology. Quat. Int. 164–165,

359 207–220 (2007).

360 49. Marom, A., McCullagh, J. S. O., Higham, T. F. G., Sinitsyn, A. A. & Hedges, R. E. M. Single

361 amino acid of Upper Paleolithic modern humans. Proc. Natl. Acad. Sci.

362 109, 6878–6881 (2012).

19

363 50. Trinkaus, E. et al. An from the Peştera cu Oase, Romania. Proc. Natl.

364 Acad. Sci. 100, 11231–11236 (2003).

365 51. Posth, C. et al. Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-

366 Africans and a Late Glacial Population Turnover in Europe. Curr. Biol. 26, 827–833 (2016).

367 52. Nalawade-Chavan, S., McCullagh, J. & Hedges, R. New Hydroxyproline Radiocarbon Dates

368 from , Russia, Confirm Early Mid Upper Palaeolithic Burials in . PLoS ONE 9,

369 e76896 (2014).

370

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Table S1. Comparative 14C dates. All the dates were calibrated using the software OxCal v4.3.2 based on the IntCal13 calibration data set1. The calibrated dates are rounded to 5. Site Individual Cultural Reference cultural 14C σ(+) σ (-) cal BP (2ρ) Laboratory Material Reference layer layer (BP) Code dated datation low high Peştera cu Oase Oase1 No direct 2 34,290 970 870 GrA-22810 Direct date 2 cultural on human association >35,20 Ox-A-11711 remains 0 900 890 41,640 37,580 Combined age 34,950 Kostenki Kostenki14 No direct 3 33,250 500 500 38,680 36,260 OxA-X-2395- Direct date 4 (Martina Gora) cultural 15 on human association remains Buran-Kaya BuranKaya3A Layer 6-1 5 31,900 240 220 36,310 35,240 GrA-37938 Dated on 5 III Gravettian human remains from the same layer Goyet Goyet Q116-1 No direct 6 30,880 170 160 35,160 34,430 GrA-46175 Direct date 6 cultural on human association remains Sunghir Sunghir3 Cultural For the whole site : 30,000 550 550 35,150 33,030 OxA-X-2395-7 Direct date 4 assignment Late Sungirian7,8 on human discussed Late Streletskian 4,9–11 remains Eastern 24,830 110 110 OxA-15754 Gravettian12,13 Kostenki 12 Kostenki12 Attributed to 14 28,500 140 140 32,995 31,840 GrA-5552 Associated 14 (Volkovskaya Gorodtsovian charcoal site) Pavlov I Pavlov1 Gravettian 15 26,170 450 450 31,100 29,420 GrN-20391 Associated 15 (evolved charcoal Pavlovian) Dolní Věstonice Vestonice16 Gravettian 15 25,740 210 210 30,570 29,390 GrN-15277 Associated 15 II (evolved charcoal Pavlovian) 25,570 280 280 30,535 29,035 GrN-15276

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Table S2. Shotgun library results

Total PCR Average % reads hg37d5 duplicates Mapping length endogenous Library >27bp Mapped reads removed Quality 20 (bp) DNA SP61-8-17_BKA4_B 6441522 26702 25125 19480 35.0 0.41% SP62-1_BKA4_B_UNG 27957579 117303 107828 85025 37.1 0.42% SP62-2_BKA4_B 10456937 75619 63145 46579 35.5 0.72% SP62-3_BKA4_B 40315968 156287 151945 117997 39.7 0.39% SP62-4_BKA4_B 38353291 151652 147546 115046 39.6 0.40% SP62-5_BKA4_B 34800662 124997 122028 95191 39.0 0.36% SP62-6_BKA4_B 32251522 109869 107635 84208 38.5 0.34% SP62-7_BKA4_B 36758191 141531 138038 107792 38.7 0.39% SP62-8_BKA4_B 20324986 108640 106323 82621 38.8 0.53% Total reads 247660658 1012600 969613 753939 38.0 0.44%

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Table S3. Y SNP summary with Palaeolithic comparisons

Y haplogroup BuranKaya3A Palaeolithic haplogroups SNPs Derived Ancestral BT 6 0 16 17 CT 1 0 Kostenki12 , 3/5 Natufians C 1 0 C1a* GoyetQ116-116 18 16 C1a2 0 7 Sunghir3 , Vestonice16 3 C1b* Kostenki14 C1b1a1 0 1 C1b1a2 0 6 C1b1a3 0 1 C1b2a 0 1 C2 0 7

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Table S4. Neanderthal content of BuranKaya3A and three ancient genomes recalculated using the BuranKaya3A subset of 6,252 Neanderthal- and modern human-derived SNPs, compared with the previously published median results calculated from either archaic ancestry or f4-ratio method from Fu et al., 201616.

BK SNP subset 95%CI Fu-Ancestry Fu-f4 BuranKaya3A 3.55% 0.0224 Oase 8.82% 0.0973 7.5% 9.9% Kostenki14 2.94% 0.0205 2.8% 3.6% Ust-Ishim 4.69% 0.0236 3.0% 4.4%

24

Table S5. Mitochondrial sequence references

Haplogroup Age* Location Culture Reference African L3 modern West Africa (Yoruba) - AF347014.1 BuranKaya3A N1 35775 Crimea Early Gravettian this study Cioclovina1 U 33212 Romania - 6 Fumane2 R 39805 - 19 Goyet2878-21 U5 26662 Belgium Gravettian 6 GoyetQ116-1 M 34795 Belgium Aurignacian 6 GoyetQ376-19 U2 27515 Belgium Gravettian 6 GoyetQ376-3 M 33540 Belgium Aurignacian 6 GoyetQ53-1 U2 27975 Belgium Gravettian 6 GoyetQ55-2 U2 27520 Belgium Gravettian 6 GoyetQ56-16 U2 26320 Belgium Gravettian 6 20 Kostenki14 U2 37470 Russia - LaRochette M 27592 France 6 Muierii1 U6 35204 Romania - 21 N1a N1a NA NA NA N1a defining mutations N1b N1b NA NA NA N1b defining mutations Natufian9 N1 10800 Isreal Natufian 17 Oase1 N 39610 Romania - 22 Paglicci108 U2'3'4'7'8'9 28396 Italy Gravettian 6 Paglicci133 U8c 33000 Italy Gravettian 6 Salkhit N 34425 Mongolia - 23 Sunghir1 U8 32820 Russia Eastern Gravettian/Streletskian 18 Sunghir2 U2 34230 Russia Eastern Gravettian/Streletskian 18 Sunghir3 U2 34090 Russia Eastern Gravettian/Streletskian 18 Sunghir4 U2 33988 Russia Eastern Gravettian/Streletskian 18 Tianyuan R 40328 China - 23 Ust-Ishim R 45020 - 24

25

Vestonice13 U8 31155 Czech Republic Gravettian 25 Vestonice14 U5 31155 Czech Republic Gravettian 25 Vestonice16 U5 29980 Czech Republic Gravettian 6 Vestonice43 U5 29977 Czech Republic Gravettian 6 *median value (calBP) from dates given in reference

26

Table S6. Genome sequence references

Age* Location Culture Reference AfontovaGora3 16710 Russia - 16 BerryAuBac 7245 France 16 Bichon 13665 Switzerland 26 Bockstein 8265 Germany Mesolithic 16 Brillenhohle 14780 Germany Magdelenian 16 BuranKaya3A 35775 Crimea Gravettian This study Burkhardtshohle 14615 Germany Magdelenian 16 Chaudardes1 8205 France Mesolithic 16 Cioclovina1 32435 Romania Mesolithic 16 Continenza 10855 Italy Mesolithic 16 ElMiron 18720 Magdelenian 16 Falkenstein 9200 Germany Mesolithic 16 GoyetQ116-1 34795 Belgium Aurignacian 16 GoyetQ-2 15005 Belgium Magdelenian 16 GoyetQ376-19 27515 Belgium Gravettian 16 GoyetQ53-1 27975 Belgium Gravettian 16 GoyetQ56-16 26320 Belgium Gravettian 16 HohleFels49 15130 Germany Magdelenian 16 HohleFels79 14670 Germany Magdelenian 16 Hotu 8878 Iran Mesolithic 17 Iboussieres39 11725 France Epipaleolithic 16 Karelia 8375 Russia Mesolithic 27 Kostenki12 32415 Russia Attributed to Gorodtsovian 16 Kostenki14 37470 Russia - 16,28 Kotias 9720 Georgia Mesolithic 26 KremsWA3 30970 Austria Gravettian 16

27

LaBrana1 7815 Spain Mesolithic 29 LesCloseaux13 9562 France Mesolithic 16 Loschbour 8050 Luxembourg Mesolithic 30 Mal'ta 24305 Russia Mal'ta - Buret' 31 Muierii2 33300 Romania - 16 Natufian9 10800 Natufian 17 Natufian6 10800 Israel Natufian 17 Oase1 39610 Romania - 22 Ofnet 8245 Germany Mesolithic 16 Ostuni1 27620 Italy Gravettian 16 Ostuni2 28975 Italy Gravettian 16 Paglicci108 28396 Italy Gravettian 16 Paglicci133 32895 Italy Gravettian 16 Pavlov1 30260 Czech Republic Gravettian 16 Ranchot88 10085 France Mesolithic 16 Rigney1 15465 France Magdelenian 16 Rochedane 12960 France Epipaleolithic 16 Satsurblia 13255 Georgia Epigravettian 26 Sunghir3 34090 Russia Eastern 18 Gravettian/Streletskian Tianyuan 40328 China - 32 Ust-Ishim 45020 Siberia - 24 Vestonice13 31155 Czech Republic Gravettian 16 Vestonice14 31155 Czech Republic Gravettian 16 Vestonice15 31155 Czech Republic Gravettian 16 Vestonice16 29980 Czech Republic Gravettian 16 Vestonice43 29977 Czech Republic Gravettian 16 Villabruna 13980 Italy Epigravettian 16 *median value (calBP) from dates given in reference

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Table S7. f3(BuranKaya3A, x, Mbuti) results

All SNPS Transversions only f3 sterr Z SNPs f3 sterr Z SNPs BuranKaya3A AfontovaGora3 Mbuti 0.384319 0.035709 10.762 1327 0.350285 0.07188 4.873 298 BuranKaya3A BerryAuBac Mbuti 0.332715 0.079569 4.181 226 0.341718 0.144269 2.369 52 BuranKaya3A Bichon Mbuti 0.402716 0.015075 26.714 8262 0.413134 0.021643 19.088 3862 BuranKaya3A Bockstein Mbuti 0.419447 0.147753 2.839 83 0.528223 0.459984 1.148 11 BuranKaya3A Brillenhohle Mbuti 0.128968 0.103485 1.246 51 0.091156 0.174642 0.522 10 BuranKaya3A Burkhardtshohle Mbuti 0.385832 0.117609 3.281 131 0.611612 0.338867 1.805 28 BuranKaya3A Chaudardes1 Mbuti 0.515568 0.083193 6.197 309 0.424476 0.173462 2.447 65 BuranKaya3A Cioclovina1 Mbuti 0.520214 0.187446 2.775 62 0.175386 0.345346 0.508 15 BuranKaya3A Continenza Mbuti 0.477686 0.263526 1.813 48 0.592877 0.81353 0.729 13 BuranKaya3A ElMiron Mbuti 0.391415 0.021648 18.081 3547 0.354761 0.035589 9.968 1066 BuranKaya3A Falkenstein Mbuti 0.348055 0.074946 4.644 231 0.401859 0.213542 1.882 37 BuranKaya3A GoyetQ116-1 Mbuti 0.402532 0.022416 17.958 3659 0.387871 0.045382 8.547 795 BuranKaya3A GoyetQ-2 Mbuti 0.453124 0.077433 5.852 263 0.605961 0.191084 3.171 61 BuranKaya3A GoyetQ376-19 Mbuti 0.276116 0.106582 2.591 88 0.223004 0.136924 1.629 17 BuranKaya3A GoyetQ53-1 Mbuti 0.185602 0.175285 1.059 34 0.323704 0.534602 0.606 6 BuranKaya3A GoyetQ56-16 Mbuti 0.553546 0.272756 2.029 28 0.352032 0.56187 0.627 7 BuranKaya3A HohleFels49 Mbuti 0.543713 0.09323 5.832 262 0.494036 0.250399 1.973 41 BuranKaya3A HohleFels79 Mbuti 0.240254 0.153623 1.564 42 0.091954 0.021719 4.234 11 BuranKaya3A Hotu Mbuti 0.376847 0.050784 7.421 691 0.261641 0.088716 2.949 153 BuranKaya3A Iboussieres39 Mbuti 0.504506 0.331982 1.52 29 0.247283 0.397081 0.623 8 BuranKaya3A Karelia Mbuti 0.422672 0.01737 24.333 7067 0.432769 0.024782 17.463 3247 BuranKaya3A Kostenki12 Mbuti 0.406626 0.075608 5.378 310 0.489396 0.174374 2.807 88 BuranKaya3A Kostenki14 Mbuti 0.430879 0.01726 24.964 7311 0.427491 0.024095 17.742 3089 BuranKaya3A Kotias Mbuti 0.393417 0.014959 26.3 8252 0.397889 0.021343 18.643 3869 BuranKaya3A KremsWA3 Mbuti 0.386445 0.039692 9.736 1031 0.527171 0.098493 5.352 231

29

BuranKaya3A LaBrana1 Mbuti 0.405692 0.015797 25.681 7271 0.414868 0.022658 18.31 3305 BuranKaya3A LesCloseaux13 Mbuti 0.156012 0.104236 1.497 35 0.156353 0.200758 0.779 5 BuranKaya3A Loschbour Mbuti 0.420236 0.014939 28.13 8324 0.425356 0.021072 20.186 3900 BuranKaya3A Mal’ta Mbuti 0.395182 0.017644 22.398 6446 0.402512 0.025462 15.808 2966 BuranKaya3A Muierii Mbuti 0.481571 0.067999 7.082 477 0.68802 0.142958 4.813 142 BuranKaya3A Natuian9 Mbuti 0.374678 0.027978 13.392 2070 0.402556 0.061442 6.552 454 BuranKaya3A Natufian6 Mbuti 0.293264 0.049968 5.869 617 0.447189 0.122606 3.647 145 BuranKaya3A Oase1 Mbuti 0.329862 0.035003 9.424 1158 0.301321 0.055523 5.427 459 BuranKaya3A Ofnet Mbuti 0.397787 0.262171 1.517 23 0.150049 0.314613 0.477 5 BuranKaya3A Ostuni1 Mbuti 0.365527 0.03162 11.56 1622 0.418664 0.060023 6.975 418 BuranKaya3A Ostuni2 Mbuti 0.45448 0.165329 2.749 83 0.307497 0.521756 0.589 15 BuranKaya3A Paglicci108 Mbuti 0.773895 0.509567 1.519 13 -0.125 1.608895 -0.078 5 BuranKaya3A Paglicci133 Mbuti 0.322502 0.069246 4.657 319 0.402489 0.164335 2.449 68 BuranKaya3A Pavlov1 Mbuti 0.449836 0.080795 5.568 299 0.478315 0.193927 2.466 65 BuranKaya3A Ranchot88 Mbuti 0.398034 0.032273 12.333 1738 0.387009 0.064378 6.011 343 BuranKaya3A Rigney1 Mbuti 0.41524 0.103769 4.002 150 0.591176 0.454989 1.299 27 BuranKaya3A Rochedane Mbuti 0.380682 0.039607 9.612 995 0.515008 0.091489 5.629 215 BuranKaya3A Satsurblia Mbuti 0.408289 0.017349 23.534 5702 0.42183 0.024122 17.487 2711 BuranKaya3A Sunghir3 Mbuti 0.448631 0.015876 28.258 8348 0.448689 0.022101 20.302 3922 BuranKaya3A Tianyuan Mbuti 0.365052 0.015869 23.004 5690 0.364851 0.023197 15.728 2558 BuranKaya3A Ust-Ishim Mbuti 0.401416 0.013894 28.891 9358 0.414576 0.019685 21.06 4375 BuranKaya3A Vestonice13 Mbuti 0.394869 0.047024 8.397 679 0.297689 0.080165 3.713 189 BuranKaya3A Vestonice14 Mbuti 0.141359 0.216404 0.653 20 0.335942 0.457878 0.734 5 BuranKaya3A Vestonice15 Mbuti 0.406163 0.119414 3.401 148 0.167227 0.193605 0.864 30 BuranKaya3A Vestonice16 Mbuti 0.435233 0.0218 19.965 4088 0.437643 0.035735 12.247 1326 BuranKaya3A Vestonice43 Mbuti 0.373642 0.048209 7.751 745 0.419048 0.08648 4.846 253 BuranKaya3A Villabruna Mbuti 0.377008 0.017081 22.072 5295 0.349925 0.026086 13.414 1852

30

References

1. Reimer, P. J. et al. INTCAL13 and MARINE13 radiocarbon age calibration curves 0–50,000 years CAL BP. Radiocarbon 55, 1869–1887

(2013).

2. Trinkaus, E. et al. An early modern human from the Peştera cu Oase, Romania. Proc. Natl. Acad. Sci. 100, 11231–11236 (2003).

3. Seguin-Orlando, A. et al. Genomic structure in Europeans dating back at least 36,200 years. Science 346, 1113–1118 (2014).

4. Marom, A., McCullagh, J. S. O., Higham, T. F. G., Sinitsyn, A. A. & Hedges, R. E. M. Single amino acid radiocarbon dating of Upper

Paleolithic modern humans. Proc. Natl. Acad. Sci. 109, 6878–6881 (2012).

5. Prat, S. et al. The Oldest Anatomically Modern Humans from Far Southeast Europe: Direct Dating, Culture and Behavior. PLoS ONE 6,

e20834 (2011).

6. Posth, C. et al. Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population

Turnover in Europe. Curr. Biol. 26, 827–833 (2016).

7. Bader, O.N. Sungir’: Verkhnepaleoliticheskaya stoyanka [Sungir: An Upper Paleolithic site]. (Nauka, 1978).

8. Vassilyev, S., Sinitsyn, A. & Marcel, O. Le Sungirien. Actes du colloque de Saint-Pétersbourg. ERAUL 147, (Etudes et Recherches

Archéologiques de l’Université de Liège, 2017).

9. Anikovich, M. V. et al. Early Upper Paleolithic in Eastern Europe and Implications for the Dispersal of Modern Humans. Science 315, 223–

226 (2007).

31

10. Bradley, B., Anikovich, M. & Engenii, G. Early Upper Paeolithic in the Russian Plain: Streletskayan flaked stone artefacts and .

Antiquity 69, 989–998 (1995).

11. Anikovich, M. Early Upper Paleolithic industries of Eastern Europe. J. World Prehistory 6, 205–245 (1992).

12. Pettitt, P. B. & Bader, N. O. Direct AMS radiocarbon dates for the Sungir mid Upper Palaeolithic burials. Antiquity 74, 269–270 (2000).

13. Dobrovolskaya, M., Richards, M.-P. & Trinkaus, E. Direct radiocarbon dates for the Mid Upper Paleolithic (eastern Gravettian) burials from

Sunghir, Russia. Bull. Mém. Société Anthropol. Paris 24, 96–102 (2012).

14. Sinitsyn, A. A. Les Sepultures de Kostenki: Chronologie, Attribution Culturelle, Rite Funéraire. in La Spiritualité, Actes du colloque de la

commission 8 de l’UISPP (Paléolithique supérieur), Liège, 10-12 décembre 2003. 237–244 (Etudes et Recherches Archeologiques de

l’Universite de Liege, 2004).

15. Trinkaus, E. & Svoboda, J. Early modern in Central Europe : the people of Dolní Věstonice and Pavlov. (Oxford University

Press., 2006).

16. Fu, Q. et al. The genetic history of Europe. 534, 200–205 (2016).

17. Lazaridis, I. et al. Genomic insights into the origin of farming in the ancient . Nature 536, 419–424 (2016).

18. Sikora, M. et al. Ancient genomes show social and reproductive behavior of early Upper Paleolithic foragers. Science 358, 659–662 (2017).

19. Benazzi, S. et al. The makers of the Protoaurignacian and implications for Neandertal extinction. Science 348, 793–796 (2015).

20. Krause, J. et al. A Complete mtDNA Genome of an Early Modern Human from Kostenki, Russia. Curr. Biol. 20, 231–236 (2010).

32

21. Hervella, M. et al. The mitogenome of a 35,000-year-old sapiens from Europe supports a Palaeolithic back-migration to Africa. Sci.

Rep. 6, (2016).

22. Fu, Q. et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature 524, 216–219 (2015).

23. Fu, Q. et al. DNA analysis of an early modern human from Tianyuan Cave, China. Proc. Natl. Acad. Sci. 110, 2223–2227 (2013).

24. Fu, Q. et al. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature 514, 445–449 (2014).

25. Fu, Q. et al. A Revised Timescale for Human Evolution Based on Ancient Mitochondrial Genomes. Curr. Biol. 23, 553–559 (2013).

26. Jones, E. R. et al. Upper Palaeolithic genomes reveal deep roots of modern Eurasians. Nat. Commun. 6, (2015).

27. Haak, W. et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207–211 (2015).

28. Seguin-Orlando, A. et al. Genomic structure in Europeans dating back at least 36,200 years. Science 346, 1113–1118 (2014).

29. Olalde, I. et al. Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. Nature 507, 225–228 (2014).

30. Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513, 409–413 (2014).

31. Raghavan, M. et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature 505, 87–91 (2014).

32. Yang, M. A. et al. 40,000-Year-Old Individual from Provides Insight into Early Population Structure in Eurasia. Curr. Biol. 27, 3202-

3208.e9 (2017).

33