Genetic admixture and language shift in the medieval Volga-Oka interfluve

[Ajeje Brazorf]

Highlights

• Iron Age inhabitants of Suzdal were genetically unique but close to Uralic speakers
• A shift in the local gene pool coincided with Slavic migrations and a language shift
• Genetic changes mirror the insights from historical linguistics and written records
• Outliers suggest far-reaching contacts during the medieval times

Summary

The Volga-Oka interfluve in northwestern Russia has an intriguing history of population influx and language shift during the Common Era. Today, most inhabitants of the region speak Russian, but until medieval times, northwestern Russia was inhabited by Uralic-speaking peoples. A gradual shift to Slavic languages started in the second half of the first millennium with the expansion of Slavic tribes, which led to the foundation of the Kievan Rus’ state in the late 9th century CE. The medieval Rus’ was multicultural and multilingual—historical records suggest that its northern regions comprised Slavic and Uralic peoples ruled by Scandinavian settlers. In the 10th–11th centuries, the introduction of Christianity and Cyrillic literature raised the prestige status of Slavic, driving a language shift from Uralic to Slavic. This eventually led to the disappearance of the Uralic languages from northwestern Russia. Here, we study a 1,500-year time transect of 30 ancient genomes and stable isotope values from the Suzdal region in the Volga-Oka interfluve. We describe a previously unsampled local Iron Age population and a gradual genetic turnover in the following centuries. Our time transect captures the population shift associated with the spread of Slavic languages and illustrates the ethnically mixed state of medieval Suzdal principality, eventually leading to the formation of the admixed but fully Slavic-speaking population that inhabits the area today. We also observe genetic outliers that highlight the importance of the Suzdal region in medieval times as a hub of long-reaching contacts via trade and warfare.

https://www.cell.com/current-biology...822(22)01826-7

Sample	Year	Cluster	Coverage	mtDNA	Y-DNA	Location
BOL002	188	VolgaOka_IA	76,12%	U4c1	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL006	192	*	70,04%	T2g	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL005	219	VolgaOka_IA	49,59%	V7a1	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL004	263	*	58,65%	U4c1	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL008	289	VolgaOka_IA	79,74%	H13a2b2a	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL003	292	VolgaOka_IA	9,86%	H5a1a	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL001	294	VolgaOka_IA	39,51%	H11a2	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL009	295	VolgaOka_IA	76,14%	H28a	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
BOL007	373	VolgaOka_IA	77,58%	H5	*	Davydovskoe Bol'shoe, Ivanovo Oblast, Russia, 155007
SHE003	841	VolgaOka_MA1	65,34%	U5a2a1b	G2a2b1a	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
GOR001	859	VolgaOka_MA1	55,44%	K1c1h	R1a1a1b1a1b1	Sovetskaya Ulitsa, 5, Gorokhovets, Vladimirskaya oblast', Russia, 601480
SHE001	930	VolgaOka_MA2	8,51%	U5a1d1	E1b1b1a1b1	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHE004	936	VolgaOka_MA1	74,09%	H6a1a4	*	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHE002	1010	*	5,35%	NA	*	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHE006	1012	VolgaOka_MA1	54,22%	K1a30a	*	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHE005	1046	VolgaOka_MA2	12,84%	HV10	R1a1a	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
GOS003	1090	VolgaOka_MA2	13,28%	I1a1	K	Sovetskaya Ulitsa, 5, Gorokhovets, Vladimirskaya oblast', Russia, 601480
GOS002	1111	VolgaOka_MA2	34,67%	U3a3	R1a1a1b1a2a	Sovetskaya Ulitsa, 5, Gorokhovets, Vladimirskaya oblast', Russia, 601480
SHE007	1125	VolgaOka_MA2	18,63%	H49	I2a2	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
GOS001	1157	VolgaOka_MA2	49,33%	I1a1a	I2a1b2a1	Sovetskaya Ulitsa, 5, Gorokhovets, Vladimirskaya oblast', Russia, 601480
SHK002	1179	SHK	28,02%	Z3c	N1	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHK001	1207	SHK	48,50%	M9a1a	J2a	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHE009	1346	*	55,64%	U2e1b1	R1a1a1b1a1	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
SHE008	1348	*	26,15%	H3	*	Gavrilovo-Posadsky District, Ivanovo Oblast, Russia
KED004	1414	*	68,70%	F2e	*	Kideksha, Vladimir Oblast, Russia, 601293
KRS001	1429	*	81,91%	T1a	*	Krasnogvardeiskii, Vladimir Oblast, Russia, 601263
KED003	1634	VolgaOka_H	80,19%	I1a1a3a	*	Kideksha, Vladimir Oblast, Russia, 601293
KBL003	1728	*	26,18%	U5b1b1a	*	Suzdalsky District, Vladimir Oblast, Russia
KED001	1747	VolgaOka_H	75,13%	H24a	*	Kideksha, Vladimir Oblast, Russia, 601293
KBL001	1756	*	NA	NA	*	Suzdalsky District, Vladimir Oblast, Russia
KED002	1772	VolgaOka_H	52,77%	H50	*	Kideksha, Vladimir Oblast, Russia, 601293
KBL002	1787	VolgaOka_H	81,75%	J1c3	*	Suzdalsky District, Vladimir Oblast, Russia

[Ajeje Brazorf]

Graphical abstract



Figure 1. Geographic locations, radiocarbon dates, and PCA and ADMIXTURE results of the 32 samples from the Volga-Oka interfluve.





Figure 3. Results from qpAdm analyses.
(A) Distal model fitted on selected ancient and present-day groups. Ancient populations are in the order from the oldest to youngest as indicated by an arrow. Present-day groups are in the descending order by Siberian ancestry. Error bars indicate one standard error. p values from chi-square test for each model are shown inside the square brackets. We show models that have a p value R 0.01. Right populations used: Ethiopia_4500BP.SG, CHG, Raqefet_M_Natufian, Onge, Villabruna, ANE, Mixe, and SHG.
(B) Sequential proximal models for the Volga-Oka time transect groups. Right populations: Ethiopia_4500BP.SG, Krasnoyarsk_Krai_BA, WEHG, EEHG,
Yamnaya_Samara, and LBK_EN. LBK_EN, European Neolithic farmers; EEHGs, East European hunter-gatherers; WEHGs, West European hunter-gatherers.



Figure S3. PCA calculated based on genetic variation of 60 West Eurasian populations. Ancient samples are projected on the principal components. Individuals included in each analysis cluster are indicated by colored outlines. Individuals excluded from analysis clusters are outlined with black. Genetic outliers discussed in detail in Results and Discussion are marked with arrows.

[Ajeje Brazorf]

Outliers highlight the connectedness of the medieval Suzdal

We detected several genetic outliers in our dataset. On the West Eurasian PCA, one individual from Bolshoye Davydovskoye 2 (BOL006) falls closer to their medieval successors than the main VolgaOka_IA group (Figure S3). Similarly, one individual from Shekshovo 9 (SHE008) falls closer to Central and West Europeans than the rest of the Shekshovo 9 individuals. Most strikingly, both individuals from Shekshovo 2 (SHK001 and SHK002) fell far from other individuals in our dataset, closer to East Asia and the ‘‘forest-steppe cline’’ in the Eurasian-wide PCA. Their closest PCA neighbors were Kazakhs, Karakalpaks, Siberian Tatars, and other Turkic-speaking groups from Central Asia and Siberia. Both individuals also carried mitochondrial haplogroups that are more common in Asia than in Europe. In cladality f4 tests, we found the smallest number of significantly non-zero estimates with Karakalpaks, suggesting genetic similarity. In a previous study, their strontium values indicated a non-local origin, further supporting the interpretation that these individuals had moved to the Volga area in their adulthood. These two men, who died at a young age, may represent Turkic-speaking groups whose members were in the military service in Kievan Rus’, mostly guarding its southern borders. Lastly, the earliest individual from Kideksha (KED004) carried an intriguing mixture of ancestries. On the West Eurasian PCA, this individual falls within the space between European and Iranian clines. This individual also carried a geographically unusual mitochondrial haplogroup F2e, mostly found in present-day East Asians and mainland Southeast Asians. ADMIXTURE suggested that this individual has genetic affinity to ancient and present-day Iranians, and thus we fitted qpAdm models with temporally proximate Iranian-related sources. The best model included medieval Alans from northern Caucasus in addition to VolgaOka_IA and Slavic-like ancestry. These findings highlight the connectedness and importance of the Volga-Oka interfluve during the medieval era.

[Ajeje Brazorf]

G25 preliminary coordinates sorted by autosomal cluster

RUS_Vladimir_1:BOL001__AD_294,0.112685,0.026404,0.084098,0.089148,0.007078,0.01255,0.016921,0.019384,-0.011453,-0.04155,0.006658,-0.012739,0.02215,0.001514,-0.009229,-0.016043,-0.010431,-0.006081,-0.008296,0.001251,0.003494,-0.001113,-0.003574,-0.022654,0.003832
RUS_Vladimir_1:BOL002__AD_188,0.112685,0.014217,0.097674,0.097546,-0.001539,0.014223,0.014806,0.014076,-0.001227,-0.035901,0.015914,-0.005095,0.014271,-0.017065,-0.00855,-0.001724,-0.000522,0.001014,-0.003017,0.001751,0.003119,0.002597,0.00419,-0.003374,-0.001317
RUS_Vladimir_1:BOL004__AD_263,0.10927,0.00914,0.098051,0.090117,-0.00954,0.013945,0.011986,0.019615,-0.001432,-0.034443,0.013153,-0.01139,0.018137,-0.007432,-0.012351,-0.008618,-0.014342,0.001647,0.000503,0.004752,0.007861,-0.004328,-0.003328,0.00241,0.002155
RUS_Vladimir_1:BOL005__AD_219,0.120652,0.036559,0.093903,0.076551,0.001231,0.013945,0.00705,0.017076,-0.00409,-0.053213,0.015265,-0.010191,0.035381,0.004129,-0.024158,0.000928,0.018515,-0.002407,-0.006034,0.006628,0.00025,-0.00371,0.012078,-0.000602,0.00012
RUS_Vladimir_1:BOL007__AD_373,0.113823,0.032497,0.089755,0.084303,0.007386,0.00753,0.00987,0.015922,-0.007567,-0.043554,0.014453,-0.012889,0.025421,-0.006881,-0.011943,-0.003713,0.004172,-0.005701,0.005531,0.002251,-0.000499,0.002844,0.002218,-0.004699,-0.000958
RUS_Vladimir_1:BOL008__AD_289,0.113823,0.019295,0.098051,0.095931,-0.001539,0.018686,0.015276,0.009692,-0.010431,-0.046835,0.016239,-0.01124,0.024975,0.008945,-0.015608,-0.000928,-0.003651,0.004561,-0.008673,0.001251,0.006114,-0.006801,0.001356,-0.015183,0.00012
RUS_Vladimir_1:BOL009__AD_295,0.117238,0.041637,0.085229,0.084626,0.000923,0.020917,0.012926,0.013615,-0.0045,-0.040274,0.011692,-0.009591,0.016799,-0.000413,-0.010586,0.013789,0.018123,-0.005068,-0.003771,-0.000625,0.007487,0.000495,0.002465,0.000241,0.005269
RUS_Vladimir_2:BOL006__AD_192,0.121791,0.066009,0.090132,0.08721,0.016926,0.026216,0.004935,0.017538,-0.0045,-0.035172,0.012342,-0.007643,0.028246,0.01913,-0.011129,0.006629,0.005346,0.001267,-0.007793,0.001501,0.003244,-0.005441,0.001725,-0.011206,-0.002515
RUS_Vladimir_2:GOR001__AD_859,0.117238,0.07718,0.082967,0.076228,0.022773,0.019243,0.013866,0.009923,0.003477,-0.035172,0.005846,-0.009591,0.018434,0.010459,-0.011401,-0.001989,0.003912,-0.006334,-0.000251,0.004502,-0.003743,-0.004575,0.0053,0.005784,-0.002515
RUS_Vladimir_2:SHE003__AD_841,0.117238,0.045699,0.086361,0.08075,0.01908,0.02008,0.012926,0.018922,-0.006749,-0.029522,0.013153,-0.016335,0.029732,0.023671,-0.013572,-0.012861,-0.008475,-0.00152,0.005531,-0.003627,0.000374,-0.001978,0.00037,-0.017593,-0.002036
RUS_Vladimir_2:SHE004__AD_936,0.120652,0.07718,0.089378,0.091086,0.022158,0.02259,0.00846,0.00923,0,-0.037541,0.009743,-0.014987,0.023042,0.012386,-0.020358,0.008751,0.01708,-0.011655,-0.006662,0.008879,0.001373,-0.005317,0.000616,-0.009158,0.000599
RUS_Vladimir_2:SHE006__AD_1012,0.114961,0.069056,0.084475,0.073967,0.017542,0.012271,0.002585,0.013384,0.000614,-0.031345,-0.001299,-0.012589,0.02334,0.009496,-0.011943,0.005569,0.020601,-0.004307,-0.000754,0.01088,-0.000374,-0.00643,-0.006286,-0.000482,0.007544
RUS_Vladimir_3:GOS001__AD_1157,0.120652,0.102568,0.078818,0.05814,0.031698,0.015618,0.010575,0.014538,0.000818,-0.012392,-0.006496,-0.01124,0.013231,0.012386,-0.003122,0.000796,0.005737,-0.001394,-0.000754,0.006378,0.005615,0.000371,0.005176,-0.003494,-0.007784
RUS_Vladimir_3:GOS002__AD_1111,0.127482,0.11577,0.075801,0.069768,0.034468,0.026774,0.011045,0.012692,0.000205,-0.021504,0.002436,-0.001199,0.015015,0.023258,-0.002714,-0.008353,-0.01343,-0.002534,0.010056,0.00075,-0.007612,-0.005317,0.007025,0.000843,-0.008263
RUS_Vladimir_3:KBL002__AD_1787,0.121791,0.1046,0.075424,0.065246,0.030775,0.023706,0.010105,0.014538,-0.001636,-0.020957,0.001461,-0.005995,0.015907,0.020231,-0.013979,0.003182,0.002738,0.00114,0.00176,0,-0.002371,-0.007914,0.011216,0.002289,0.001796
RUS_Vladimir_3:KBL003__AD_1728,0.119514,0.093429,0.067882,0.058786,0.039392,0.018128,-0.00846,0.005769,-0.010226,-0.022962,-0.000812,-0.004946,0.012636,0.025323,-0.010179,-0.00305,0.003651,-0.003927,0.010182,0.001376,-0.006239,-0.001978,0.004314,-0.003615,-0.008622
RUS_Vladimir_3:KED001__AD_1747,0.127482,0.107646,0.07731,0.069122,0.025235,0.023985,0.00846,0.021461,-0.0045,-0.03098,0.002436,-0.009442,0.020367,0.02546,-0.005157,-0.009414,-0.012647,-0.002027,-0.003645,0.00075,-0.002745,0.004451,0.006655,0.000723,0.001916
RUS_Vladimir_3:KED002__AD_1772,0.129758,0.105615,0.07995,0.06783,0.036314,0.016455,0.00705,0.012692,-0.005113,-0.024784,0.002436,-0.007643,0.019475,0.014313,-0.00475,0.009546,0.004824,0.000127,0.005028,-0.006378,0,0.001113,0.018117,0.001205,0.009221
RUS_Vladimir_3:KED003__AD_1634,0.133173,0.105615,0.07731,0.063308,0.031698,0.01506,0.013631,0.016384,0.001841,-0.023326,0.005846,-0.013638,0.010704,0.018579,-0.012893,-0.003447,0.000522,-0.009122,0.006913,0.007504,-0.005116,-0.010634,0.001972,-0.009519,-0.006945
RUS_Vladimir_3:KRS001__AD_1429,0.126344,0.097491,0.075047,0.068476,0.021542,0.024821,0.009165,0.010615,-0.00225,-0.018406,-0.001624,-0.009891,0.021258,0.013487,-0.016286,0.00769,0.019036,-0.005068,-0.00088,0.003126,0.009358,0.002844,0.00419,0.005784,-0.008263
RUS_Vladimir_3:SHE005__AD_1046,0.110408,0.121864,0.075801,0.073644,0.023697,0.034025,0.002585,0.010153,-0.002454,-0.036447,0.00065,-0.007343,0.016501,0.025185,-0.006515,-0.003182,-0.004303,-0.007601,0.002263,-0.004627,0.000873,-0.012984,0.005053,-0.011929,-0.004431
RUS_Vladimir_3:SHE007__AD_1125,0.113823,0.113739,0.066373,0.059109,0.01908,0.012829,0.008225,0.018922,0.000205,-0.018041,-0.003248,-0.005395,0.022745,0.032204,-0.011129,-0.010077,-0.005867,-0.004181,-0.003771,0.010255,0.013102,-0.008285,0.005793,-0.011688,-0.005389
RUS_Vladimir_3:SHE009__AD_1346,0.119514,0.107646,0.065242,0.052326,0.039392,0.026216,0.011281,0.011769,0.000205,-0.015308,-0.008444,-0.007044,0.015609,0.024634,-0.013572,-0.002387,0.005215,-0.004054,0.013324,0.003752,0.004492,-0.009521,0.00037,0.00012,-0.001078
RUS_Vladimir_4:KED004__AD_1414,0.119514,0.094444,0.042238,0.033592,0.008617,0.008925,0.003995,0.008077,-0.016771,-0.011481,-0.000162,-0.002548,0.000595,0.012111,-0.00855,-0.012463,-0.009648,0.00114,0.007039,-0.009505,0.001747,-0.004451,0.00037,-0.006145,-0.001317
RUS_Vladimir_5:GOS003__AD_1090,0.122929,0.122879,0.04978,0.052972,0.042469,0.019243,-0.00094,0.017538,0.006954,-0.016583,0.00341,-0.005545,-0.00223,0.015138,-0.004479,0.016441,0.017863,-0.008488,-0.003142,0.001126,0.001622,-0.002597,0.016638,-0.011929,0.010418
RUS_Vladimir_6:SHE008__AD_1348,0.121791,0.122879,0.055437,0.04199,0.035083,0.023427,0.008695,0.003461,0.002659,0.007107,-0.004547,0.005395,-0.009366,-0.013487,0.016286,0.022275,0.014212,-0.003801,0.005279,0.007629,0.00861,-0.005812,-0.001849,0.011929,-0.003712
RUS_Vladimir_7:SHK001__AD_1207,0.073985,-0.182795,0.062979,0.014858,-0.04924,-0.018686,0.001645,0.00923,-0.012271,-0.006378,-0.013803,-0.002847,0.003717,-0.015138,0.002307,-0.006895,-0.009909,0.0019,0.01257,0.009379,-0.022959,0.001978,-0.00493,-0.006989,-0.002395
RUS_Vladimir_7:SHK002__AD_1179,0.066017,-0.174671,0.042615,-0.000646,-0.038469,-0.003347,0.007755,0.014769,-0.009613,-0.008383,-0.021922,-0.001499,-0.000297,-0.004817,0.002986,0.011403,-0.00326,0.003294,0.010936,0.012006,-0.018467,-0.003586,-0.016885,0.002048,-0.000599

Averages

RUS_Vladimir_1,0.11431086,0.025678429,0.092394429,0.088317429,0.00057142857,0.014542286,0.012690714,0.015625714,-0.0058142857,-0.042252857,0.013339143,-0.010447857,0.022447714,-0.0024575714,-0.013203571,-0.0023298571,0.0016948571,-0.0017192857,-0.0033938571,0.0024655714,0.0039751429,-0.0014308571,0.0022007143,-0.0062658571,0.0013172857
RUS_Vladimir_2,0.118376,0.0670248,0.0866626,0.0818482,0.0196958,0.02008,0.0085544,0.0137994,-0.0014316,-0.0337504,0.007957,-0.012229,0.0245588,0.0150284,-0.0136806,0.0012198,0.0076928,-0.0045098,-0.0019858,0.004427,0.0001748,-0.0047482,0.000345,-0.006531,0.0002154
RUS_Vladimir_3,0.12272191,0.10690755,0.074087091,0.064159545,0.030299182,0.021601545,0.0076056364,0.013593909,-0.0020822727,-0.022282455,-0.00048718182,-0.007616,0.016677091,0.021369091,-0.0091178182,-0.0016996364,0.00049781818,-0.0035128182,0.0036796364,0.0020805455,0.00085063636,-0.0043503636,0.0063528182,-0.0026619091,-0.0034401818
RUS_Vladimir_4,0.119514,0.094444,0.042238,0.033592,0.008617,0.008925,0.003995,0.008077,-0.016771,-0.011481,-0.000162,-0.002548,0.000595,0.012111,-0.00855,-0.012463,-0.009648,0.00114,0.007039,-0.009505,0.001747,-0.004451,0.00037,-0.006145,-0.001317
RUS_Vladimir_5,0.122929,0.122879,0.04978,0.052972,0.042469,0.019243,-0.00094,0.017538,0.006954,-0.016583,0.00341,-0.005545,-0.00223,0.015138,-0.004479,0.016441,0.017863,-0.008488,-0.003142,0.001126,0.001622,-0.002597,0.016638,-0.011929,0.010418
RUS_Vladimir_6,0.121791,0.122879,0.055437,0.04199,0.035083,0.023427,0.008695,0.003461,0.002659,0.007107,-0.004547,0.005395,-0.009366,-0.013487,0.016286,0.022275,0.014212,-0.003801,0.005279,0.007629,0.00861,-0.005812,-0.001849,0.011929,-0.003712
RUS_Vladimir_7,0.070001,-0.178733,0.052797,0.007106,-0.0438545,-0.0110165,0.0047,0.0119995,-0.010942,-0.0073805,-0.0178625,-0.002173,0.00171,-0.0099775,0.0026465,0.002254,-0.0065845,0.002597,0.011753,0.0106925,-0.020713,-0.000804,-0.0109075,-0.0024705,-0.001497

Distances

Distance to:	RUS_Vladimir_1
0.02716834	Russian_Leshukonsky
0.02993012	Saami_Kola
0.02998968	Komi
0.04850213	Russian_Pinezhsky
0.05196850	Russian_Pinega

Distance to:	RUS_Vladimir_2
0.02204465	Vepsian
0.02221318	Russian_Pinezhsky
0.02252796	Russian_Pinega
0.02462559	Russian_Krasnoborsky
0.02675214	Karelian

Distance to:	RUS_Vladimir_3
0.01358258	Russian_Ryazan
0.01450500	Russian_Tver
0.01605313	Russian_Kursk
0.01662030	Russian_Orel
0.01873338	Ukrainian_Dnipro

Distance to:	RUS_Vladimir_4
0.05006917	Moldovan
0.05012817	Turkish_Deliorman
0.05543867	Ukrainian_Zakarpattia
0.05550358	Moldovan_o
0.05666444	Hungarian

Distance to:	RUS_Vladimir_5
0.03861691	Slovakian
0.04005217	Moldovan_o
0.04177744	Ukrainian_Zakarpattia
0.04241052	Czech
0.04499683	Sorb_Niederlausitz

Distance to:	RUS_Vladimir_6
0.03477232	Afrikaner
0.03511915	Dutch
0.03859357	French_Brittany
0.03869940	French_Pas-de-Calais
0.03876305	German

Distance to:	RUS_Vladimir_7
0.02891882	Karakalpak
0.03415786	Nogai
0.04939685	Kazakh
0.05073268	Tubalar
0.06179607	Tatar_Siberian

G25 preliminary coordinates sorted by period

RUS_Vladimir_AD_1-500:BOL001__AD_294,0.112685,0.026404,0.084098,0.089148,0.007078,0.01255,0.016921,0.019384,-0.011453,-0.04155,0.006658,-0.012739,0.02215,0.001514,-0.009229,-0.016043,-0.010431,-0.006081,-0.008296,0.001251,0.003494,-0.001113,-0.003574,-0.022654,0.003832
RUS_Vladimir_AD_1-500:BOL002__AD_188,0.112685,0.014217,0.097674,0.097546,-0.001539,0.014223,0.014806,0.014076,-0.001227,-0.035901,0.015914,-0.005095,0.014271,-0.017065,-0.00855,-0.001724,-0.000522,0.001014,-0.003017,0.001751,0.003119,0.002597,0.00419,-0.003374,-0.001317
RUS_Vladimir_AD_1-500:BOL004__AD_263,0.10927,0.00914,0.098051,0.090117,-0.00954,0.013945,0.011986,0.019615,-0.001432,-0.034443,0.013153,-0.01139,0.018137,-0.007432,-0.012351,-0.008618,-0.014342,0.001647,0.000503,0.004752,0.007861,-0.004328,-0.003328,0.00241,0.002155
RUS_Vladimir_AD_1-500:BOL005__AD_219,0.120652,0.036559,0.093903,0.076551,0.001231,0.013945,0.00705,0.017076,-0.00409,-0.053213,0.015265,-0.010191,0.035381,0.004129,-0.024158,0.000928,0.018515,-0.002407,-0.006034,0.006628,0.00025,-0.00371,0.012078,-0.000602,0.00012
RUS_Vladimir_AD_1-500:BOL006__AD_192,0.121791,0.066009,0.090132,0.08721,0.016926,0.026216,0.004935,0.017538,-0.0045,-0.035172,0.012342,-0.007643,0.028246,0.01913,-0.011129,0.006629,0.005346,0.001267,-0.007793,0.001501,0.003244,-0.005441,0.001725,-0.011206,-0.002515
RUS_Vladimir_AD_1-500:BOL007__AD_373,0.113823,0.032497,0.089755,0.084303,0.007386,0.00753,0.00987,0.015922,-0.007567,-0.043554,0.014453,-0.012889,0.025421,-0.006881,-0.011943,-0.003713,0.004172,-0.005701,0.005531,0.002251,-0.000499,0.002844,0.002218,-0.004699,-0.000958
RUS_Vladimir_AD_1-500:BOL008__AD_289,0.113823,0.019295,0.098051,0.095931,-0.001539,0.018686,0.015276,0.009692,-0.010431,-0.046835,0.016239,-0.01124,0.024975,0.008945,-0.015608,-0.000928,-0.003651,0.004561,-0.008673,0.001251,0.006114,-0.006801,0.001356,-0.015183,0.00012
RUS_Vladimir_AD_1-500:BOL009__AD_295,0.117238,0.041637,0.085229,0.084626,0.000923,0.020917,0.012926,0.013615,-0.0045,-0.040274,0.011692,-0.009591,0.016799,-0.000413,-0.010586,0.013789,0.018123,-0.005068,-0.003771,-0.000625,0.007487,0.000495,0.002465,0.000241,0.005269
RUS_Vladimir_AD_500-1000:GOR001__AD_859,0.117238,0.07718,0.082967,0.076228,0.022773,0.019243,0.013866,0.009923,0.003477,-0.035172,0.005846,-0.009591,0.018434,0.010459,-0.011401,-0.001989,0.003912,-0.006334,-0.000251,0.004502,-0.003743,-0.004575,0.0053,0.005784,-0.002515
RUS_Vladimir_AD_500-1000:SHE003__AD_841,0.117238,0.045699,0.086361,0.08075,0.01908,0.02008,0.012926,0.018922,-0.006749,-0.029522,0.013153,-0.016335,0.029732,0.023671,-0.013572,-0.012861,-0.008475,-0.00152,0.005531,-0.003627,0.000374,-0.001978,0.00037,-0.017593,-0.002036
RUS_Vladimir_AD_500-1000:SHE004__AD_936,0.120652,0.07718,0.089378,0.091086,0.022158,0.02259,0.00846,0.00923,0,-0.037541,0.009743,-0.014987,0.023042,0.012386,-0.020358,0.008751,0.01708,-0.011655,-0.006662,0.008879,0.001373,-0.005317,0.000616,-0.009158,0.000599
RUS_Vladimir_AD_1000-1500:GOS001__AD_1157,0.120652,0.102568,0.078818,0.05814,0.031698,0.015618,0.010575,0.014538,0.000818,-0.012392,-0.006496,-0.01124,0.013231,0.012386,-0.003122,0.000796,0.005737,-0.001394,-0.000754,0.006378,0.005615,0.000371,0.005176,-0.003494,-0.007784
RUS_Vladimir_AD_1000-1500:GOS002__AD_1111,0.127482,0.11577,0.075801,0.069768,0.034468,0.026774,0.011045,0.012692,0.000205,-0.021504,0.002436,-0.001199,0.015015,0.023258,-0.002714,-0.008353,-0.01343,-0.002534,0.010056,0.00075,-0.007612,-0.005317,0.007025,0.000843,-0.008263
RUS_Vladimir_AD_1000-1500:KRS001__AD_1429,0.126344,0.097491,0.075047,0.068476,0.021542,0.024821,0.009165,0.010615,-0.00225,-0.018406,-0.001624,-0.009891,0.021258,0.013487,-0.016286,0.00769,0.019036,-0.005068,-0.00088,0.003126,0.009358,0.002844,0.00419,0.005784,-0.008263
RUS_Vladimir_AD_1000-1500:SHE005__AD_1046,0.110408,0.121864,0.075801,0.073644,0.023697,0.034025,0.002585,0.010153,-0.002454,-0.036447,0.00065,-0.007343,0.016501,0.025185,-0.006515,-0.003182,-0.004303,-0.007601,0.002263,-0.004627,0.000873,-0.012984,0.005053,-0.011929,-0.004431
RUS_Vladimir_AD_1000-1500:SHE006__AD_1012,0.114961,0.069056,0.084475,0.073967,0.017542,0.012271,0.002585,0.013384,0.000614,-0.031345,-0.001299,-0.012589,0.02334,0.009496,-0.011943,0.005569,0.020601,-0.004307,-0.000754,0.01088,-0.000374,-0.00643,-0.006286,-0.000482,0.007544
RUS_Vladimir_AD_1000-1500:SHE007__AD_1125,0.113823,0.113739,0.066373,0.059109,0.01908,0.012829,0.008225,0.018922,0.000205,-0.018041,-0.003248,-0.005395,0.022745,0.032204,-0.011129,-0.010077,-0.005867,-0.004181,-0.003771,0.010255,0.013102,-0.008285,0.005793,-0.011688,-0.005389
RUS_Vladimir_AD_1000-1500:SHE009__AD_1346,0.119514,0.107646,0.065242,0.052326,0.039392,0.026216,0.011281,0.011769,0.000205,-0.015308,-0.008444,-0.007044,0.015609,0.024634,-0.013572,-0.002387,0.005215,-0.004054,0.013324,0.003752,0.004492,-0.009521,0.00037,0.00012,-0.001078
RUS_Vladimir_AD_1500-2000:KBL002__AD_1787,0.121791,0.1046,0.075424,0.065246,0.030775,0.023706,0.010105,0.014538,-0.001636,-0.020957,0.001461,-0.005995,0.015907,0.020231,-0.013979,0.003182,0.002738,0.00114,0.00176,0,-0.002371,-0.007914,0.011216,0.002289,0.001796
RUS_Vladimir_AD_1500-2000:KBL003__AD_1728,0.119514,0.093429,0.067882,0.058786,0.039392,0.018128,-0.00846,0.005769,-0.010226,-0.022962,-0.000812,-0.004946,0.012636,0.025323,-0.010179,-0.00305,0.003651,-0.003927,0.010182,0.001376,-0.006239,-0.001978,0.004314,-0.003615,-0.008622
RUS_Vladimir_AD_1500-2000:KED001__AD_1747,0.127482,0.107646,0.07731,0.069122,0.025235,0.023985,0.00846,0.021461,-0.0045,-0.03098,0.002436,-0.009442,0.020367,0.02546,-0.005157,-0.009414,-0.012647,-0.002027,-0.003645,0.00075,-0.002745,0.004451,0.006655,0.000723,0.001916
RUS_Vladimir_AD_1500-2000:KED002__AD_1772,0.129758,0.105615,0.07995,0.06783,0.036314,0.016455,0.00705,0.012692,-0.005113,-0.024784,0.002436,-0.007643,0.019475,0.014313,-0.00475,0.009546,0.004824,0.000127,0.005028,-0.006378,0,0.001113,0.018117,0.001205,0.009221
RUS_Vladimir_AD_1500-2000:KED003__AD_1634,0.133173,0.105615,0.07731,0.063308,0.031698,0.01506,0.013631,0.016384,0.001841,-0.023326,0.005846,-0.013638,0.010704,0.018579,-0.012893,-0.003447,0.000522,-0.009122,0.006913,0.007504,-0.005116,-0.010634,0.001972,-0.009519,-0.006945
RUS_Vladimir_o1_AD_1000-1500:KED004__AD_1414,0.119514,0.094444,0.042238,0.033592,0.008617,0.008925,0.003995,0.008077,-0.016771,-0.011481,-0.000162,-0.002548,0.000595,0.012111,-0.00855,-0.012463,-0.009648,0.00114,0.007039,-0.009505,0.001747,-0.004451,0.00037,-0.006145,-0.001317
RUS_Vladimir_o2_AD_1000-1500:GOS003__AD_1090,0.122929,0.122879,0.04978,0.052972,0.042469,0.019243,-0.00094,0.017538,0.006954,-0.016583,0.00341,-0.005545,-0.00223,0.015138,-0.004479,0.016441,0.017863,-0.008488,-0.003142,0.001126,0.001622,-0.002597,0.016638,-0.011929,0.010418
RUS_Vladimir_o3_AD_1000-1500:SHE008__AD_1348,0.121791,0.122879,0.055437,0.04199,0.035083,0.023427,0.008695,0.003461,0.002659,0.007107,-0.004547,0.005395,-0.009366,-0.013487,0.016286,0.022275,0.014212,-0.003801,0.005279,0.007629,0.00861,-0.005812,-0.001849,0.011929,-0.003712
RUS_Vladimir_o4_AD_1000-1500:SHK001__AD_1207,0.073985,-0.182795,0.062979,0.014858,-0.04924,-0.018686,0.001645,0.00923,-0.012271,-0.006378,-0.013803,-0.002847,0.003717,-0.015138,0.002307,-0.006895,-0.009909,0.0019,0.01257,0.009379,-0.022959,0.001978,-0.00493,-0.006989,-0.002395
RUS_Vladimir_o4_AD_1000-1500:SHK002__AD_1179,0.066017,-0.174671,0.042615,-0.000646,-0.038469,-0.003347,0.007755,0.014769,-0.009613,-0.008383,-0.021922,-0.001499,-0.000297,-0.004817,0.002986,0.011403,-0.00326,0.003294,0.010936,0.012006,-0.018467,-0.003586,-0.016885,0.002048,-0.000599

Averages

RUS_Vladimir_AD_1-500,0.11524587,0.03071975,0.092111625,0.088179,0.00261575,0.0160015,0.01172125,0.01586475,-0.00565,-0.04136775,0.0132145,-0.01009725,0.0231725,0.000240875,-0.01294425,-0.00121,0.00215125,-0.001346,-0.00394375,0.002345,0.00388375,-0.001932125,0.00214125,-0.006883375,0.00083825
RUS_Vladimir_AD_500-1000,0.118376,0.066686333,0.086235333,0.082688,0.021337,0.020637667,0.011750667,0.012691667,-0.0010906667,-0.034078333,0.0095806667,-0.013637667,0.023736,0.015505333,-0.015110333,-0.002033,0.0041723333,-0.006503,-0.00046066667,0.0032513333,-0.00066533333,-0.0039566667,0.0020953333,-0.006989,-0.0013173333
RUS_Vladimir_AD_1000-1500,0.11902629,0.10401914,0.074508143,0.065061429,0.026774143,0.021793429,0.007923,0.013153286,-0.00037957143,-0.021920429,-0.002575,-0.0078144286,0.018242714,0.020092857,-0.0093258571,-0.0014205714,0.0038555714,-0.0041627143,0.0027834286,0.0043591429,0.0036362857,-0.0056174286,0.0030458571,-0.002978,-0.003952
RUS_Vladimir_AD_1500-2000,0.1263436,0.103381,0.0755752,0.0648584,0.0326828,0.0194668,0.0061572,0.0141688,-0.0039268,-0.0246018,0.0022734,-0.0083328,0.0158178,0.0207812,-0.0093916,-0.0006366,-0.0001824,-0.0027618,0.0040476,0.0006504,-0.0032942,-0.0029924,0.0084548,-0.0017834,-0.0005268
RUS_Vladimir_o1_AD_1000-1500,0.119514,0.094444,0.042238,0.033592,0.008617,0.008925,0.003995,0.008077,-0.016771,-0.011481,-0.000162,-0.002548,0.000595,0.012111,-0.00855,-0.012463,-0.009648,0.00114,0.007039,-0.009505,0.001747,-0.004451,0.00037,-0.006145,-0.001317
RUS_Vladimir_o2_AD_1000-1500,0.122929,0.122879,0.04978,0.052972,0.042469,0.019243,-0.00094,0.017538,0.006954,-0.016583,0.00341,-0.005545,-0.00223,0.015138,-0.004479,0.016441,0.017863,-0.008488,-0.003142,0.001126,0.001622,-0.002597,0.016638,-0.011929,0.010418
RUS_Vladimir_o3_AD_1000-1500,0.121791,0.122879,0.055437,0.04199,0.035083,0.023427,0.008695,0.003461,0.002659,0.007107,-0.004547,0.005395,-0.009366,-0.013487,0.016286,0.022275,0.014212,-0.003801,0.005279,0.007629,0.00861,-0.005812,-0.001849,0.011929,-0.003712
RUS_Vladimir_o4_AD_1000-1500,0.070001,-0.178733,0.052797,0.007106,-0.0438545,-0.0110165,0.0047,0.0119995,-0.010942,-0.0073805,-0.0178625,-0.002173,0.00171,-0.0099775,0.0026465,0.002254,-0.0065845,0.002597,0.011753,0.0106925,-0.020713,-0.000804,-0.0109075,-0.0024705,-0.001497

Distances

Distance to:	RUS_Vladimir_AD_1-500
0.02276799	Russian_Leshukonsky
0.03009342	Komi
0.03200895	Saami_Kola
0.04321339	Russian_Pinezhsky
0.04627169	Russian_Pinega

Distance to:	RUS_Vladimir_AD_500-1000
0.02159230	Russian_Pinega
0.02256984	Vepsian
0.02279975	Russian_Pinezhsky
0.02480987	Russian_Krasnoborsky
0.02738269	Karelian

Distance to:	RUS_Vladimir_AD_1000-1500
0.01823023	Russian_Ryazan
0.01886823	Russian_Tver
0.02188903	Russian_Kursk
0.02384382	Russian_Orel
0.02403307	Moksha

Distance to:	RUS_Vladimir_AD_1500-2000
0.01268650	Russian_Tver
0.01437833	Russian_Ryazan
0.01765745	Russian_Kursk
0.01793742	Russian_Orel
0.01983446	Ukrainian_Dnipro

Distance to:	RUS_Vladimir_o1_AD_1000-1500
0.05006917	Moldovan
0.05012817	Turkish_Deliorman
0.05543867	Ukrainian_Zakarpattia
0.05550358	Moldovan_o
0.05666444	Hungarian

Distance to:	RUS_Vladimir_o2_AD_1000-1500
0.03861691	Slovakian
0.04005217	Moldovan_o
0.04177744	Ukrainian_Zakarpattia
0.04241052	Czech
0.04499683	Sorb_Niederlausitz

Distance to:	RUS_Vladimir_o3_AD_1000-1500
0.03477232	Afrikaner
0.03511915	Dutch
0.03859357	French_Brittany
0.03869940	French_Pas-de-Calais
0.03876305	German

Distance to:	RUS_Vladimir_o4_AD_1000-1500
0.02891882	Karakalpak
0.03415786	Nogai
0.04939685	Kazakh
0.05073268	Tubalar
0.06179607	Tatar_Siberian

[rothaer]

Graphical abstract



Figure 1. Geographic locations, radiocarbon dates, and PCA and ADMIXTURE results of the 32 samples from the Volga-Oka interfluve.

I wonder what caused the genetic change from the second to the third stage. An out-of-proportion reproduction of Slavs before mixing?

[Ajeje Brazorf]

I wonder what caused the genetic change from the second to the third stage. An out-of-proportion reproduction of Slavs before mixing?

Change was gradual and it's quite likely that Slavs simply outnumbered the pre-Slavic communities.

[rothaer]

Change was gradual and it's quite likely that Slavs simply outnumbered the pre-Slavic communities.

The pic "Middle Ages" after the "Slavic migrations" does, however, not yet show such an outnumbering. So if from that pic it has developed without any additional notable Slavic migration to such an outnumbering - I also assume such before the mixing that resulted in the pic "Post-medieval" - then this outnumbering must have emerged out of a very differing reproduction rate. Or how to explain?

[Russki]

Distance to: RUS_Vladimir_o3_AD_1000-1500
0.03477232 Afrikaner
0.03511915 Dutch
0.03859357 French_Brittany
0.03869940 French_Pas-de-Calais
0.03876305 German

German spy

[sevruk]

I wonder what caused the genetic change from the second to the third stage. An out-of-proportion reproduction of Slavs before mixing?

Slavs have more dick than the Uralics

[sevruk]

VolgaOka_MA2 is hardly suitable even for pure Eastern Slavs, rather something Central Russian/Moksha/Erzya