Genetic study The Genetic Origin of the Indo-Europeans

[Archetype0ne]

The Genetic Origin of the Indo-Europeans

The Yamnaya archaeological complex appeared around 3300BCE across the steppes north of the Black and Caspian Seas, and by 3000BCE reached its maximal extent from Hungary in the west to Kazakhstan in the east. To localize the ancestral and geographical origins of the Yamnaya among the diverse...

www.biorxiv.org

"

Abstract​

The Yamnaya archaeological complex appeared around 3300BCE across the steppes north of the Black and Caspian Seas, and by 3000BCE reached its maximal extent from Hungary in the west to Kazakhstan in the east. To localize the ancestral and geographical origins of the Yamnaya among the diverse Eneolithic people that preceded them, we studied ancient DNA data from 428 individuals of which 299 are reported for the first time, demonstrating three previously unknown Eneolithic genetic clines. First, a "Caucasus-Lower Volga" (CLV) Cline suffused with Caucasus hunter-gatherer (CHG) ancestry extended between a Caucasus Neolithic southern end in Neolithic Armenia, and a steppe northern end in Berezhnovka in the Lower Volga. Bidirectional gene flow across the CLV cline created admixed intermediate populations in both the north Caucasus, such as the Maikop people, and on the steppe, such as those at the site of Remontnoye north of the Manych depression. CLV people also helped form two major riverine clines by admixing with distinct groups of European hunter-gatherers. A "Volga Cline" was formed as Lower Volga people mixed with upriver populations that had more Eastern hunter-gatherer (EHG) ancestry, creating genetically hyper-variable populations as at Khvalynsk in the Middle Volga. A "Dnipro Cline" was formed as CLV people bearing both Caucasus Neolithic and Lower Volga ancestry moved west and acquired Ukraine Neolithic hunter-gatherer (UNHG) ancestry to establish the population of the Serednii Stih culture from which the direct ancestors of the Yamnaya themselves were formed around 4000BCE. This population grew rapidly after 3750-3350BCE, precipitating the expansion of people of the Yamnaya culture who totally displaced previous groups on the Volga and further east, while admixing with more sedentary groups in the west. CLV cline people with Lower Volga ancestry contributed four fifths of the ancestry of the Yamnaya, but also, entering Anatolia from the east, contributed at least a tenth of the ancestry of Bronze Age Central Anatolians, where the Hittite language, related to the Indo-European languages spread by the Yamnaya, was spoken. We thus propose that the final unity of the speakers of the "Proto-Indo-Anatolian" ancestral language of both Anatolian and Indo-European languages can be traced to CLV cline people sometime between 4400-4000 BCE."

 

[Archetype0ne]

23abcThe sample you've all been waiting for came out in the 2nd IE preprint released today.

I10206, Moldova_EBA_Yamnaya, 2900-2500 BCE, J-L283

Grave 12:10 (individual ID I10206)

The grave was situated in the northeastern sector of the mound, 2.80 m away from the central landmark (R.C.), at a depth of 1.65 m. The burial chamber, rectangular with rounded corners and slightly arched sides, was oriented east-west, measuring 1.25 × 1.80 m. Based on the characteristics of the filling of the pit, the arrangement of skeletal remains, and the location of the grave goods, it can be inferred that grave no.10/M.10 was disturbed in ancient times. The skeletal remains were found scattered in different areas of the pit and at different depths.

It was deduced that the deceased was originally placed at the bottom of the pit, with the head oriented to the east. Traces from the plant layer covering the pit bottom were found near and beneath the skeleton. Grave goods: a flint arrowhead of elongated triangular shape with slightly convex sides and a concave base. The artefacts’ length is 2 cm, width 1.4 cm, and thickness 0.5cm.

The skeleton, lacking anatomical connection, is incomplete and poorly preserved. Traces of bright red ochre were identified on all its elements, particularly intense on the skull. Other taphonomic changes observed include cracking, exfoliation (aerial weathering), and marks left by rodent teeth.

Anthropological sex: male.
Molecular sex: male, with an unusual Y-haplogroup J2b2a1 (J-L283).
Biological age at death: about 30 years (young adult).
Pathologies: supragingival calculus; active porotic hyperostosis (cribra cranii).

The skeleton is very robust, with extremely pronounced muscle insertions, with extensive enthesopathic changes on the humerus and femurs. The individual exhibits a very large skeletal stature, at least 181 cm.

Traumas: a perimortem fracture in the middle third of the right clavicle.

The burial must be attributed to Cernavodă-I or Yamnaya cultures.

 

[Archetype0ne]

"Pre-Indo-Anatolian languages were spoken by at least some of these diverse ancestors, but living Indo-European languages trace their ongins to the Yamnaya expansion ca. 3300BCE out of (3) and the earlier Indo-Anatolian expansion ca 4400-4000BCE out of (1)
Dripro"

 

[Archetype0ne]

Archetype0neSome figures I found interesting:

Ps: Guess everything is interesting.

Admixture table:

 

[Maciamo]

Important paper. Over 350 samples. The supplementary material PDF even has facial reconstructions by archaeological culture!

Here is a summary of the Y-DNA and mtDNA.

First the reference Neolithic samples.

Master ID	Date (BP)	Group ID for analysis	Country	mtDNA	Y-DNA	Y-DNA
I1685	12700	Natufian	Israel	n/a (<2x)	E-Z830	E1b1b
I1690	12700	Natufian	Israel	n/a (<2x)	..	..
I7527	9900	Iran_GanjDareh_N	Iran	U7a	n/a (female)	n/a (female)
I8554	9900	Levant_N	Jordan	n/a (<2x)	CT	CT
I1945	9800	Iran_GanjDareh_N	Iran	J1	R-Y3399	R2
I1710	9565	Levant_N	Jordan	T1a2	E-Z1919	E1b1b1a1
I1699	8700	Levant_N	Jordan	R0a2	n/a (female)	n/a (female)
I6910	8341	Labazy	Russia	U4a1	Q-M930	Q1b1a
I1099	8300	TUR_Marmara_Barcın_N	Turkey	T2b	G-Z6484	G2a2a1a2
I1100	8300	TUR_Marmara_Barcın_N	Turkey	K1a	n/a (female)	n/a (female)
I1102	8300	TUR_Marmara_Barcın_N	Turkey	K1a3a	C-V86	C1a2
I1103	8300	TUR_Marmara_Barcın_N	Turkey	K1b1b1	G-Z6484	G2a2a1a2
I0726	7950	TUR_Marmara_Menteşe_N	Turkey	H	n/a (female)	n/a (female)
I3931	7858	Armenia_Aknashen_N	Armenia	I1	J-PF5174	J2a1a1b1a1a
I1671	7686	Iran_LN_SehGabi	Iran	K1a12a	G-Z6553	G2a
I2532	7614	Romania_EN	Romania	K1a2	G-PF3359	G2a2b2b
I2529	7610	Bulgaria_N	Bulgaria	T1a	I-M436	I2a1b
I3930	7535	Armenia_MasisBlur_N	Armenia	K1a12a1a	J-Y13341	J2a1a1a2a

Then the Steppe samples sorted by culture group.

Master ID	Date (BP)	Group ID for analysis	Country	mtDNA	Y-DNA	Y-DNA
I22199	6780	BPgroup	Russia	H13a1	I-S12195	I2a1b1a2a2a2
I22200	6718	BPgroup	Russia	H2a1	R-Y106006	R1b1a2b
I22201	6331	BPgroup	Russia	U2e1'2'3	R-V1636	R1b1a2
I22202	6087	BPgroup	Russia	U5a1f	n/a (female)	n/a (female)
I6303	5528	ChekalinoIV	Russia	U5a2+16294	n/a (female)	n/a (female)
I5124	6205	Csongrád_I5124	Hungary	K1b2	Q-Y6802	Q1b
I3546	7363	Ekaterinovka	Russia	U5a1+@16192	Q-L53	Q1b1
I8740	7353	Ekaterinovka	Russia	U5a1+@16192	R-V1636	R1b1a2
I6064	7349	Ekaterinovka	Russia	U4a3	R-L754	R1b
I8287	7306	Ekaterinovka	Russia	U5a1	Q-M1107	Q1b1a
I6061	7299	Ekaterinovka	Russia	U4a3	n/a (female)	n/a (female)
I6069	7286	Ekaterinovka	Russia	U5a1+@16192	n/a (female)	n/a (female)
I8286	7264	Ekaterinovka	Russia	U4a1	Q-M930	Q1b1a
I6057	7263	Ekaterinovka	Russia	U4a2c	n/a (female)	n/a (female)
I6068	7230	Ekaterinovka	Russia	U5a1+@16192	R-L754	R1b
I6062	7216	Ekaterinovka	Russia	U5a1i	R-V1636	R1b1a2
I8282	7215	Ekaterinovka	Russia	U5a1	Q-M930	Q1b1a
I6059	7213	Ekaterinovka	Russia	U2e1f1	n/a (female)	n/a (female)
I6066	7212	Ekaterinovka	Russia	U5a1+@16192	R-V1636	R1b1a2
I6101	7210	Ekaterinovka	Russia	U4a1	n/a (female)	n/a (female)
I8283	7210	Ekaterinovka	Russia	U4d	R-L754	R1b
I8290	7207	Ekaterinovka	Russia	U5a1b	n/a (female)	n/a (female)
I23648	7200	Ekaterinovka	Russia	U5a1+@16192	n/a (female)	n/a (female)
I23649	7200	Ekaterinovka	Russia	U5a2	n/a (female)	n/a (female)
I23650	7200	Ekaterinovka	Russia	U5a1+@16192	n/a (female)	n/a (female)
I23651	7200	Ekaterinovka	Russia	U4a	Q-M930	Q1b1a
I23652	7200	Ekaterinovka	Russia	U4a2	n/a (female)	n/a (female)
I20189	7184	Ekaterinovka	Russia	U4a2	n/a (female)	n/a (female)
I8284	7113	Ekaterinovka	Russia	U5a1+@16192	n/a (female)	n/a (female)
I20192	7098	Ekaterinovka	Russia	U4a2	n/a (female)	n/a (female)
I20115	7056	Ekaterinovka	Russia	U4	n/a (female)	n/a (female)
I8285	7216	Ekaterinovka_o	Russia	U2e1b	R-V1636	R1b1a2
I20114	7200	Ekaterinovka_o	Russia	U4a1	P-P337	P1
I12492	7450	GK1	Russia	T2c1a	R-M459	R1a
I12493	7450	GK1	Russia	U2e1b	K	K
I12491	7425	GK1	Russia	U5a2b	R-M459	R1a
I12490	7450	GK2	Russia	U4b1	R-M459	R1a
I2363	5168	Hungary_EBA_Yamnaya	Hungary	U5a1b1	n/a (female)	n/a (female)
I5120	4839	Hungary_EBA_Yamnaya	Hungary	R1b1	n/a (female)	n/a (female)
I5126	4833	Hungary_EBA_Yamnaya	Hungary	H2a1a	n/a (female)	n/a (female)
I2772	4781	Hungary_EBA_Yamnaya	Hungary	U4a	n/a (female)	n/a (female)
I2761	4675	Hungary_EBA_Yamnaya	Hungary	T2g	R-Z2110	R1b1a1b1b3
I3525	4650	Hungary_EBA_Yamnaya_1drel.I3510_contam	Hungary	H2b	R-Y132510	R1b1a1b1a1a2c1
I18100	4709	Hungary_EBA_Yamnaya_o	Hungary	V	G-Z42565	G2a2a1a2b
I3511	4574	Hungary_EBA_Yamnaya_o	Hungary	V7	R-M12149	R1b1a1b1b
I2762	4847	Hungary_EBA_Yamnaya_onjp	Hungary	U2e1h	R-Z2108	R1b1a1b1b3
I18099	4720	Hungary_EBA_Yamnaya_onjp	Hungary	U5a1a1	n/a (female)	n/a (female)
I3510	4669	Hungary_EBA_Yamnaya_onjp	Hungary	H2b	R-M269	R1b1a1b
I2362	4927	Hungary_LateC_EBA_Yamnaya	Hungary	T2a1b1a	n/a (female)	n/a (female)
I18101	4725	Hungary_LateC_EBA_Yamnaya	Hungary	I2	R-M12149	R1b1a1b1b
I26302	4900	Kazakhstan_EBA_Yamnaya	Kazakhstan	W	Q-BZ1466	Q1b2b1b2b~
I26304	4900	Kazakhstan_EBA_Yamnaya	Kazakhstan	n/a (<2x)	n/a (female)	n/a (female)
I26224	4900	Kazakhstan_EBA_Yamnaya	Kazakhstan	n/a (<2x)	n/a (female)	n/a (female)
I26289	4900	Kazakhstan_EBA_Yamnaya_possible	Kazakhstan	n/a (<2x)	n/a (female)	n/a (female)
I26227	4900	Kazakhstan_Kumsay_EBA	Kazakhstan	U2e1	n/a (female)	n/a (female)
I26231	4900	Kazakhstan_Kumsay_EBA	Kazakhstan	C5c+16234	Q-L939	Q1b2b1b2b~
I26235	4900	Kazakhstan_Kumsay_EBA	Kazakhstan	H6a1b	n/a (female)	n/a (female)
I26230	4900	Kazakhstan_Kumsay_EBA_Yamnaya	Kazakhstan	U2e1b	R-Y106006	R1b1a2b
I26303	4900	Kazakhstan_Kumsay_EBA_Yamnaya	Kazakhstan	U2e1b	R-V1636	R1b1a2
I6741	6820	Khi	Russia	U4b1	R-L389	R1b1a
I6105	6665	Khi	Russia	U2e1b	n/a (female)	n/a (female)
I6737	6625	Khi	Russia	U5a1a2	R-L754	R1b
I6412	6556	Khi	Russia	H13a2a	R-L754	R1b
I6734	6525	Khi	Russia	U2e1b	n/a (female)	n/a (female)
I6739	6415	Khi	Russia	U2e2a1	Q-YP1669	Q1a2b~
I6104	6393	Khi	Russia	U2e1a1	R-L389	R1b1a
I11837	6375	Khi	Russia	U4a1c	R-L754	R1b
I6300	7076	KhlopkovBugor	Russia	U5a1f1	n/a (female)	n/a (female)
I6301	7070	KhlopkovBugor	Russia	U5a1a1	n/a (female)	n/a (female)
I6905	7042	KhlopkovBugor_contam	Russia	H13a1a1	n/a (female)	n/a (female)
I6404	7199	Klo	Russia	U5a1	n/a (female)	n/a (female)
I6110	7058	Klo	Russia	U4	n/a (female)	n/a (female)
I6740	6971	Klo	Russia	..	Q-YP1669	Q1a2b~
I6108	6951	Klo	Russia	U4a	n/a (female)	n/a (female)
I6106	6808	Klo	Russia	T2a1b	n/a (female)	n/a (female)
I6109	6716	Klo	Russia	U5a1+@16192	R-L389	R1b1a
I20188	6697	Klo	Russia	R1b	Q-YP1669	Q1a2b~
I6102	6576	Klo	Russia	T2a1b	n/a (female)	n/a (female)
I6735	6981	Kmed	Russia	U4a	J-CTS1026	J1
I6736	6837	Kmed	Russia	U5a1a1	R-L754	R1b
I6103	6831	Kmed	Russia	U5a2d	I-L699	I2a1b1a2a2a
I6107	6794	Kmed	Russia	U5a2d	R-L389	R1b1a
I6405	6782	Kmed	Russia	U5a2d	R-L389	R1b1a
I6406	6775	Kmed	Russia	U4d	R-L754	R1b
I6738	6748	Kmed	Russia	R1b1	R-V1636	R1b1a2
I0426	6375	Kmed	Russia	U4a	n/a (female)	n/a (female)
I11828	6257	Krivyansky	Russia	T2a1b	J-M319	J2a1a1a2b1b
I4429	5300	Maikop_I4429	Russia	T2a1b1a1	J-PF5119	J2a1a1a2
I6904	5964	Maximovka	Russia	U5a1d2b	I-S21825	I2a1a1b
I7847	4911	Moldova_EBA_Yamnaya	Moldova	U5a1b1	R-M12149	R1b1a1b1b
I10398	4850	Moldova_EBA_Yamnaya	Moldova	I4a	n/a (female)	n/a (female)
I7848	4806	Moldova_EBA_Yamnaya	Moldova	U4c2a	n/a (female)	n/a (female)
I10414	4693	Moldova_EBA_Yamnaya	Moldova	H6a1a	n/a (female)	n/a (female)
I10415	4653	Moldova_EBA_Yamnaya	Moldova	U4a	R-M269	R1b1a1b
I10206	4650	Moldova_EBA_Yamnaya	Moldova	U4b3	J-L283	J2b2a1
I10208	4650	Moldova_EBA_Yamnaya	Moldova	H47	R-Z2108	R1b1a1b1b3
I10424	4650	Moldova_EBA_Yamnaya	Moldova	U5a1a1+16362	n/a (female)	n/a (female)
I10174	4577	Moldova_EBA_Yamnaya	Moldova	..	R-M269	R1b1a1b
I7857	5685	Moldova_Eneolithic	Moldova	U5b1	n/a (female)	n/a (female)
I28683	5913	Remontnoye	Russia	R0	R-V1636	R1b1a2
I28682	5632	Remontnoye	Russia	H2a	n/a (female)	n/a (female)
I14119	4885	Romania_Brailita_EBA_Yamnaya	Romania	U5a1a1	I-Y5669	I2a1b1a2a2a2
I12825	4525	Romania_EBA_Catacomb	Romania	T1a1	R-M269	R1b1a1b
I12824	4490	Romania_EBA_Catacomb	Romania	U4c2	n/a (female)	n/a (female)
I10478	4850	Romania_EBA_Yamnaya	Romania	U5a1d	K	K
I12823	4850	Romania_EBA_Yamnaya	Romania	K1b2b	R-L51	R1b1a1b1a
I10499	4745	Romania_EBA_Yamnaya	Romania	T2c1a	R-Z2108	R1b1a1b1b3
I10500	4736	Romania_EBA_Yamnaya	Romania	H2c	R-PF7562	R1b1a1b
I10498	4725	Romania_EBA_Yamnaya	Romania	U4a	n/a (female)	n/a (female)
I10486	4703	Romania_EBA_Yamnaya	Romania	H6a1a	R1b	R1b
I10891	4690	Romania_EBA_Yamnaya	Romania	U4c1	R-M12149	R1b1a1b1b
I18820	4669	Romania_EBA_Yamnaya	Romania	H2a1	n/a (female)	n/a (female)
I18818	4715	Romania_EBA_Yamnaya_1d.rel.I18819_contam	Romania	K1c1	P-P337	P1
I18819	4700	Romania_EBA_Yamnaya_oSouth	Romania	H2a	Q-M25	Q1a2
I2533	7371	Romania_EN	Romania	J1c5	n/a (female)	n/a (female)
I26089	5082	Russia_Afanasievo	Russia	H6a1b	R-Z2108	R1b1a1b1b3
I5269	4902	Russia_Afanasievo	Russia	T2a1a	R-Z2108	R1b1a1b1b3
I13813	4850	Russia_Afanasievo	Russia	T2d1a	R-Z2108	R1b1a1b1b3
I13812	4850	Russia_Afanasievo	Russia	U5a1a1	R-Z2108	R1b1a1b1b3
I25159	4650	Russia_Afanasievo	Russia	W3a1	n/a (female)	n/a (female)
I25160	4650	Russia_Afanasievo	Russia	U5a2+16294	n/a (female)	n/a (female)
I25161	4650	Russia_Afanasievo	Russia	U2e1	n/a (female)	n/a (female)
I3388	4650	Russia_Afanasievo	Russia	U5a1d2b	n/a (female)	n/a (female)
I20101	4897	Russia_Afanasievo_contam	Russia	T2a1b1a1b	R-Z2108	R1b1a1b1b3
I20090	4800	Russia_Afanasievo_contam	Russia	R1b	R-M269	R1b1a1b
I20541	4661	Russia_Afanasievo_Khakassia_possible	Russia	T2c1a2	Q-Y6826	Q1b2a1a~
I20089	4685	Russia_Afanasievo_oOkunevo	Russia	H6a1b	N-Y6503	N
I20559	4937	Russia_Afanasievo_Yenisei	Russia	U4a1	R-Y13200	R1b1a1a
I29570	4500	Russia_BA_ECatacomb	Russia	T2a1b1a	R-M269	R1b1a1b
RISE662	4067	Russia_BA_Okunevo	Russia	H6a1b	Q-Y18330	Q1b1a3a1~
I29573	4300	Russia_BA_WestManych_Catacomb	Russia	U4a1	n/a (female)	n/a (female)
I33307	5577	Russia_CaspianInland_EBA_Yamnaya	Russia	U4a1	R-Z2106	R1b1a1b1b3
I33863	5075	Russia_CaspianInland_EBA_Yamnaya	Russia	n/a (<2x)	F	F
I32873	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	T1a1	R-M12149	R1b1a1b1b
I32851	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	U5a1a1	R-KMS67	R1b1a1b1b3
I32868	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	R1a1a	R-Z2108	R1b1a1b1b3
I32998	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	n/a (<2x)	R-M269	R1b1a1b
I32864	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	n/a (<2x)	R-Z2106	R1b1a1b1b3
I32866	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	U5a1a	n/a (female)	n/a (female)
I32867	4900	Russia_CaspianInland_EBA_Yamnaya	Russia	U2e1b	n/a (female)	n/a (female)
I32865	4885	Russia_CaspianInland_EBA_Yamnaya	Russia	U5a1g1	R-Z2106	R1b1a1b1b3
I32871	4862	Russia_CaspianInland_EBA_Yamnaya	Russia	U5a1	R-KMS67	R1b1a1b1b3
I33864	4850	Russia_CaspianInland_EBA_Yamnaya	Russia	H2a+152 16311	R-L23	R1b1a1b1
I33309	4850	Russia_CaspianInland_EBA_Yamnaya	Russia	H15a1a1	n/a (female)	n/a (female)
I33867	4850	Russia_CaspianInland_EBA_Yamnaya	Russia	n/a (<2x)	n/a (sex unknown)	n/a (sex unknown)
I33868	4850	Russia_CaspianInland_EBA_Yamnaya	Russia	I4a	n/a (female)	n/a (female)
I33306	4850	Russia_CaspianInland_EBA_Yamnaya	Russia	U5a1a1	R-M269	R1b1a1b
I32996	4804	Russia_CaspianInland_EBA_Yamnaya	Russia	U3c	R-M269	R1b1a1b
I32872	4775	Russia_CaspianInland_EBA_Yamnaya	Russia	I1b	R-M12149	R1b1a1b1b
I32876	4775	Russia_CaspianInland_EBA_Yamnaya	Russia	W3a1a	n/a (female)	n/a (female)
I33308	4740	Russia_CaspianInland_EBA_Yamnaya	Russia	H6a1b	R-FT91192	R1b1a1b1b3
I10567	4715	Russia_CaspianInland_EBA_Yamnaya	Russia	U4a1e	R-KMS67	R1b1a1b1b3
I6885	4700	Russia_CaspianInland_EBA_Yamnaya	Russia	U4	R-Z2108	R1b1a1b1b3
I32999	4681	Russia_CaspianInland_EBA_Yamnaya	Russia	U5b2a1a+16311	R-M12149	R1b1a1b1b
I32875	4653	Russia_CaspianInland_EBA_Yamnaya	Russia	U4d3	R-Z2108	R1b1a1b1b3
I33312	4400	Russia_CaspianInland_EBA_Yamnaya	Russia	U4d	R-Z2106	R1b1a1b1b3
I33315	4400	Russia_CaspianInland_EBA_Yamnaya	Russia	J1b1a1	R-L23	R1b1a1b1
I32850	4392	Russia_CaspianInland_EBA_Yamnaya	Russia	U5a1i	R-L23	R1b1a1b1
I33314	4400	Russia_CaspianInland_EBA_Yamnaya_lc	Russia	n/a (<2x)	n/a (sex unknown)	n/a (sex unknown)
I32852	4775	Russia_CaspianInland_EBA_Yamnaya_o2	Russia	K1a28	n/a (female)	n/a (female)
I7487	4726	Russia_Chelyabinsk_EBA_Yamnaya	Russia	N1a1a1a1	R-Z2108	R1b1a1b1b3
I7488	4725	Russia_Chelyabinsk_EBA_Yamnaya	Russia	T1a1	R-KMS67	R1b1a1b1b3
I7483	4721	Russia_Chelyabinsk_EBA_Yamnaya	Russia	J1b1a1	R-KMS67	R1b1a1b1b3
I7484	4721	Russia_Chelyabinsk_EBA_Yamnaya	Russia	T2a1a	n/a (female)	n/a (female)
I7486	4709	Russia_Chelyabinsk_EBA_Yamnaya	Russia	W3a1a	R-KMS67	R1b1a1b1b3
I2809	4279	Russia_Dagestan_EMBA	Russia	H13a1a1	R-M269	R1b1a1b
I12686	5175	Russia_Don_EBA_Yamnaya	Russia	U4a1	I-L699	I2a1b1a2a2a
I26778	4948	Russia_Don_EBA_Yamnaya	Russia	U5a1f2	I-L699	I2a1b1a2a2a
I24093	4902	Russia_Don_EBA_Yamnaya	Russia	U4a1e	R-L754	R1b
I24091	4891	Russia_Don_EBA_Yamnaya	Russia	U5a1g1	I-L699	I2a1b1a2a2a
I24086	4879	Russia_Don_EBA_Yamnaya	Russia	R1a1a	I-L699	I2a1b1a2a2a
I12685	4851	Russia_Don_EBA_Yamnaya	Russia	T2a1b1	I-L699	I2a1b1a2a2a
I24089	4851	Russia_Don_EBA_Yamnaya	Russia	U4a2	I-L699	I2a1b1a2a2a
I24090	4851	Russia_Don_EBA_Yamnaya	Russia	H15a1	I-L699	I2a1b1a2a2a
I6882	4850	Russia_Don_EBA_Yamnaya	Russia	K1a19a	..	..
I24088	4848	Russia_Don_EBA_Yamnaya	Russia	T2a1a	I-L699	I2a1b1a2a2a
I26787	4845	Russia_Don_EBA_Yamnaya	Russia	H26	I-L699	I2a1b1a2a2a
I26785	4842	Russia_Don_EBA_Yamnaya	Russia	U5a1j	R-M12149	R1b1a1b1b
I26780	4785	Russia_Don_EBA_Yamnaya	Russia	H2a1e	n/a (female)	n/a (female)
I12687	4775	Russia_Don_EBA_Yamnaya	Russia	T2a1a	I-L699	I2a1b1a2a2a
I8952	4750	Russia_Don_EBA_Yamnaya	Russia	U5a1a1	R-L754	R1b
I11029	4745	Russia_Don_EBA_Yamnaya	Russia	U2e1h	n/a (female)	n/a (female)
I26638	4731	Russia_Don_EBA_Yamnaya	Russia	W3a1	R-M12149	R1b1a1b1b
I10627	4722	Russia_Don_EBA_Yamnaya	Russia	U5a1a1	R-L23	R1b1a1b1
I24085	4715	Russia_Don_EBA_Yamnaya	Russia	U5a1a1	I-L699	I2a1b1a2a2a
I10628	4681	Russia_Don_EBA_Yamnaya	Russia	J2b1a1	R-L23	R1b1a1b1
I8951	4681	Russia_Don_EBA_Yamnaya	Russia	U4a1	I-L699	I2a1b1a2a2a
I26782	4644	Russia_Don_EBA_Yamnaya	Russia	U4b3	n/a (female)	n/a (female)
I26783	4596	Russia_Don_EBA_Yamnaya	Russia	U5a2d	I-L699	I2a1b1a2a2a
I12686	5175	Russia_Don_EBA_Yamnaya_A	Russia	U4a1	I-L699	I2a1b1a2a2a
I24087	4891	Russia_Don_EBA_Yamnaya_A	Russia	U5a2b	n/a (female)	n/a (female)
I8950	4794	Russia_Don_EBA_Yamnaya_A	Russia	H41a	R-M269	R1b1a1b
I6884	4614	Russia_Don_EBA_Yamnaya_A	Russia	T1a1	R-L51	R1b1a1b1a
I26781	4497	Russia_Don_EBA_Yamnaya_B	Russia	U4b1b2	n/a (female)	n/a (female)
I8949	4324	Russia_Don_EBA_Yamnaya_B	Russia	T2a1a	R-M269	R1b1a1b
I11829	5082	Russia_Don_EBA_Yamnaya_C	Russia	U3a2	n/a (female)	n/a (female)
I8477	4711	Russia_Don_EBA_Yamnaya_lc	Russia	n/a (<2x)	n/a (female)	n/a (female)
I23654	4844	Russia_Don_EMBA_Yamnaya_Catacomb_transitional	Russia	H2a1	I-L699	I2a1b1a2a2a
I23655	4757	Russia_Don_EMBA_Yamnaya_Catacomb_transitional	Russia	U5a2	I-L699	I2a1b1a2a2a
I26435	4834	Russia_EBA_Srubnaya_o1	Russia	U2e2a	R-M12149	R1b1a1b1b
I23568	4674	Russia_EBA_Yamnaya_Don	Russia	U5a2	I-L699	I2a1b1a2a2a
I6063	7293	Russia_Ekaterinovka_Eneolithic	Russia	U5a1	n/a (female)	n/a (female)
I6060	7271	Russia_Ekaterinovka_Eneolithic	Russia	U4a1	n/a (female)	n/a (female)
I8289	7236	Russia_Ekaterinovka_Eneolithic	Russia	U5a1+@16192	n/a (female)	n/a (female)
I6065	7215	Russia_Ekaterinovka_Eneolithic	Russia	U5a1+@16192	R-V1636	R1b1a2
I8288	7214	Russia_Ekaterinovka_Eneolithic	Russia	U5a1+@16192	R-V1636	R1b1a2
I6067	7212	Russia_Ekaterinovka_Eneolithic	Russia	U4a3	R-V1636	R1b1a2
I20118	7200	Russia_Ekaterinovka_Eneolithic	Russia	U5a1+@16192	n/a (female)	n/a (female)
I20116	7200	Russia_Ekaterinovka_Eneolithic	Russia	n/a (<2x)	R-P297	R1b1a1
I8738	7200	Russia_Ekaterinovka_Eneolithic	Russia	n/a (<2x)	..	..
I6058	7078	Russia_Ekaterinovka_Eneolithic	Russia	U4a3	R-V1636	R1b1a2
I8451	6448	Russia_Eneolithic	Russia	U5a1d2	Q-F1096	Q1a
I8744	6400	Russia_Eneolithic	Russia	U4a1	Q-F1096	Q1a
I11030	6348	Russia_Eneolithic	Russia	U2e2a	Q-F1096	Q1a
I8447	5820	Russia_Eneolithic	Russia	U4a1	n/a (female)	n/a (female)
I8742	5617	Russia_Eneolithic_1d.rel.I6904	Russia	U5a1d2b	I-S21825	I2a1a1b
TIM008	4473	Russia_Ivanovo_Fatyanovo_BA	Russia	K1	R-Z93	R1a1a1
I4000	4344	Russia_Kalmykia_EasternManych_EMBA	Russia	U4c1	R-M269	R1b1a1b
I3787	4316	Russia_Kalmykia_EasternManych_EMBA	Russia	U4c1	R-M269	R1b1a1b
I4004	4188	Russia_Kalmykia_EasternManych_EMBA	Russia	T1a1	BT	BT
I12883	4269	Russia_Kalmykia_EBA	Russia	T	R-M269	R1b1a1b
I12893	4900	Russia_Kalmykia_EBA_Yamnaya	Russia	H13a1a	R-L51	R1b1a1b1a
I6302	7081	Russia_KhlopkovBugor_Eneolithic	Russia	H2a1	F	F
I6408	7058	Russia_Khvalynsk_Eneolithic	Russia	U4a1	R-L754	R1b
I6299	7028	Russia_Khvalynsk_Eneolithic	Russia	U2e2a1	Q-YP1669	Q1a2b~
I6407	6831	Russia_Khvalynsk_Eneolithic	Russia	U2e1b	Q-YP1669	Q1a2b~
I6403	6820	Russia_Khvalynsk_Eneolithic	Russia	H2a	R-L754	R1b
I6402	6657	Russia_Khvalynsk_Eneolithic	Russia	H2a1	R-L754	R1b
I20298	6100	Russia_Kulmetovskiy_N	Russia	R1b	R-BY95094	R1b1a1b2b
I20890	6100	Russia_Kulmetovskiy_N	Russia	n/a (<2x)	n/a (female)	n/a (female)
I32997	4668	Russia_LowerDon_EBA_Yamnaya	Russia	T2c1a2	R-Z2106	R1b1a1b1b3
I33000	4850	Russia_LowerVolga_EBA_Yamnaya	Russia	n/a (<2x)	n/a (sex unknown)	n/a (sex unknown)
I26200	4300	Russia_MBA_Catacomb_lc	Russia	U2e1a1	BT	BT
I10632	5350	Russia_N_BA_possible	Russia	I4a	N-Z4762	N
I6297	4915	Russia_Orlovka_EBA_Yamnaya	Russia	U5b2	n/a (female)	n/a (female)
I29559	4750	Russia_Remontnoye_EBA_Yamnaya	Russia	T1a1'3	R-M12149	R1b1a1b1b
RISE547	4713	Russia_Remontnoye_EBA_Yamnaya	Russia	T2a1a	R-M12149	R1b1a1b1b
RISE240	4705	Russia_Remontnoye_EBA_Yamnaya	Russia	U5a1d1	n/a (female)	n/a (female)
RISE546	4700	Russia_Remontnoye_EBA_Yamnaya	Russia	U5a1d2b	R-Z2106	R1b1a1b1b3
I29568	4700	Russia_Remontnoye_EBA_Yamnaya	Russia	T2a1a	n/a (female)	n/a (female)
I29558	4675	Russia_Remontnoye_EBA_Yamnaya	Russia	n/a (<2x)	R-P297	R1b1a1
I29565	4500	Russia_Rostov_LateYamnaya_BA	Russia	D4m2a	n/a (female)	n/a (female)
RISE548	4600	Russia_Rostov_Steppe_NorthCaucasus_BA	Russia	U4	R-Z2106	R1b1a1b1b3
I6733	5163	Russia_Samara_EBA_Yamnaya	Russia	U5a1a1	R-M12149	R1b1a1b1b
I1450	5153	Russia_Samara_EBA_Yamnaya	Russia	W6	R-Z2103	R1b1a1b1
I0243	5052	Russia_Samara_EBA_Yamnaya	Russia	H2b	R-Z2103	R1b1a1b1
I6729	5042	Russia_Samara_EBA_Yamnaya	Russia	U5b2a1a+16311	R-Z2103	R1b1a1b1
I0242	4965	Russia_Samara_EBA_Yamnaya	Russia	H2b	n/a (female)	n/a (female)
I7490	4945	Russia_Samara_EBA_Yamnaya	Russia	J1b1a1	n/a (female)	n/a (female)
I6731	4932	Russia_Samara_EBA_Yamnaya	Russia	H13a2b2a	R-Y13200	R1b1a1a
I6728	4932	Russia_Samara_EBA_Yamnaya	Russia	T1a1	R-KMS67	R1b1a1b1b3
I6730	4930	Russia_Samara_EBA_Yamnaya	Russia	T2a1b1a1	n/a (female)	n/a (female)
I6907	4915	Russia_Samara_EBA_Yamnaya	Russia	H13a1a2	n/a (female)	n/a (female)
I10714	4850	Russia_Samara_EBA_Yamnaya	Russia	I3a	R-KMS67	R1b1a1b1b3
I7677	4844	Russia_Samara_EBA_Yamnaya	Russia	H13a2b2a	R-Z2108	R1b1a1b1b3
I25026	4830	Russia_Samara_EBA_Yamnaya	Russia	H13a2b2a	R-Z2103	R1b1a1b1
I7479	4775	Russia_Samara_EBA_Yamnaya	Russia	H13a1a1	n/a (female)	n/a (female)
I0245	4705	Russia_Samara_EBA_Yamnaya	Russia	T1a1	R-KMS67	R1b1a1b1b3
I0441	4701	Russia_Samara_EBA_Yamnaya	Russia	H2b	n/a (female)	n/a (female)
I8291	4653	Russia_Samara_EBA_Yamnaya	Russia	U5b2a1a+16311	R-Y20993	R1b1a1b1b3
I6705	4602	Russia_Samara_EBA_Yamnaya	Russia	U2e1b	R-M12149	R1b1a1b1b
I6727	4518	Russia_Samara_EBA_Yamnaya	Russia	R1a1a	R-Y87909	R1b1a1b1b3
I6906	5164	Russia_Samara_EBA_Yamnaya_possible	Russia	U4a	n/a (female)	n/a (female)
I6703	4602	Russia_StavropolKrai_EBA_Yamnaya	Russia	N1a1a1a1	R-Z2106	R1b1a1b1b3
I32619	5000	Russia_UpperOb_Eneolithic_Afanasievo	Russia	U5a1i	R-M12149	R1b1a1b1b
I32502	5000	Russia_UpperOb_Eneolithic_Afanasievo	Russia	U4	n/a (female)	n/a (female)
I33646	4950	Russia_UpperOb_Eneolithic_Afanasievo	Russia	U5a1a1	R-L23	R1b1a1b1
I32819	4950	Russia_UpperOb_Eneolithic_Afanasievo	Russia	J2a2a	R-Z2108	R1b1a1b1b3
I32822	4950	Russia_UpperOb_Eneolithic_Afanasievo	Russia	U2e1	n/a (female)	n/a (female)
I32821	4950	Russia_UpperOb_Eneolithic_Afanasievo	Russia	T2c1a2	n/a (female)	n/a (female)
I32499	4750	Russia_UpperYenisey_Eneolithic_Afanasievo	Russia	U5a1d2b	n/a (female)	n/a (female)
I32500	4750	Russia_UpperYenisey_Eneolithic_Afanasievo	Russia	U2e1h	n/a (female)	n/a (female)
I32501	4750	Russia_UpperYenisey_Eneolithic_Afanasievo	Russia	J2a2a	R-Z2106	R1b1a1b1b3
I32497	4750	Russia_UpperYenisey_Eneolithic_Afanasievo	Russia	H15b1	I-M423	I2a1a2
I11831	4918	Russia_Ural_EBA_Yamnaya	Russia	U4a1e	R-KMS67	R1b1a1b1b3
I10033	4758	Russia_Ural_EBA_Yamnaya	Russia	T1a1	n/a (female)	n/a (female)
I10034	4711	Russia_Ural_EBA_Yamnaya	Russia	U5a1b	R-Z2103	R1b1a1b1
I11990	4681	Russia_Ural_EBA_Yamnaya	Russia	U4a	R-Y87909	R1b1a1b1b3
I10630	4674	Russia_Ural_EBA_Yamnaya	Russia	U4b1b1	R-KMS67	R1b1a1b1b3
I12242	4625	Russia_Ural_EBA_Yamnaya	Russia	U5a1a1	R-Y87909	R1b1a1b1b3
I12243	4621	Russia_Ural_EBA_Yamnaya	Russia	H6a1a	R-KMS67	R1b1a1b1b3
I12241	4652	Russia_Ural_EBA_Yamnaya_contam	Russia	H2a1	R-M269	R1b1a1b
I10363	5165	Russia_Volga_EBA_Yamnaya	Russia	..	n/a (female)	n/a (female)
I11840	4848	Russia_Volga_EBA_Yamnaya	Russia	U4c1	R-Z2108	R1b1a1b1b3
I20158	4842	Russia_Volga_EBA_Yamnaya	Russia	n/a (<2x)	R-M269	R1b1a1b
I10362	4701	Russia_Volga_EBA_Yamnaya	Russia	..	n/a (female)	n/a (female)
I11838	4602	Russia_Volga_EBA_Yamnaya	Russia	U5a1a1	R-L51	R1b1a1b1a
I10441	4922	Russia_Volgograd_EBA_Yamnaya	Russia	..	R-M269	R1b1a1b
I22197	4871	Russia_Volgograd_EBA_Yamnaya	Russia	U4a2	R-Z2103	R1b1a1b1
I10597	4850	Russia_Volgograd_EBA_Yamnaya	Russia	H2a1	R-M12149	R1b1a1b1b
I6919	4596	Russia_Volgograd_EBA_Yamnaya	Russia	..	n/a (female)	n/a (female)
I6918	4564	Russia_Volgograd_EBA_Yamnaya	Russia	..	R-Z2108	R1b1a1b1b3
I6920	4850	Russia_Volgograd_EBA_Yamnaya_o	Russia	..	R-M269	R1b1a1b
NIK008AB	4508	Russia_Yaroslavl_Fatyanovo_BA	Russia	H5b	R-Z93	R1a1a1
I20499	4710	Serbia_EBA_Yamnaya	Serbia	U4b1b1	R-L52	R1b1a1b1a1
I7890	4875	Slovakia_EBA_Yamnaya	Slovakia	T1a1	R-M12149	R1b1a1b1b
I7891	4518	Slovakia_EBA_Yamnaya	Slovakia	H	R-M12149	R1b1a1b1b
I7889	4750	Slovakia_Yamnaya_lc	Slovakia	n/a (<2x)	..	..
I6559	6238	SShi	Ukraine	U4b	n/a (female)	n/a (female)
I4118	6864	SSmed	Ukraine	K1b2b	n/a (female)	n/a (female)
I5894	6561	SSmed	Ukraine	I4a	I-L699	I2a1b1a2a2a
I6558	5456	SSmed	Ukraine	U5a1f1	I-Y3259	I2a1b1
I22205	7425	Syezzheye	Russia	U2e1	n/a (female)	n/a (female)
I22204	7100	Syezzheye	Russia	R1b1	R-V1636	R1b1a2
I22203	6741	Syezzheye	Russia	U5a1d	R-V1274	R1b1a2a
I5882	5005	Ukraine_EBA_Deriivka_I5882	Ukraine	U5a2+16294	n/a (female)	n/a (female)
I5884	4735	Ukraine_EBA_Deriivka_I5884	Ukraine	U5a2b	R-M12149	R1b1a1b1b
I7840	6348	Ukraine_Eneolithic	Ukraine	n/a (<2x)	..	..
I6563	4028	Ukraine_MBA	Ukraine	U4	n/a (female)	n/a (female)
I4112	7359	Ukraine_N	Ukraine	U5a2+16294	R-V88	R1b
I3720	7233	Ukraine_N	Ukraine	U2e1h	I2	I2
I5886	7212	Ukraine_N	Ukraine	U4a	I-L701	I2a1b1a2
I5881	7077	Ukraine_N	Ukraine	U5a1	R-V2219	R1b
I3721	6951	Ukraine_N	Ukraine	U5a2+16294	n/a (female)	n/a (female)
I5891	7297	Ukraine_N_1d.rel.I4114	Ukraine	U4	R1b	R1b
I5893	7249	Ukraine_N_1d.rel.I5881	Ukraine	U5a2+16294	R-L754	R1b
I3719	6831	Ukraine_N_Deriivka_I3719	Ukraine	H1	I-CTS10057	I2a1b1a2
I5888	7349	Ukraine_N_father.or.son.I5875	Ukraine	U5b2b1	I-Y5606	I2a1b1a2a2~
I5878	7139	Ukraine_N_father.or.son.I5883	Ukraine	U4b1a1a	R-V88	R1b
Ukraine_N1	7300	Ukraine_N_son.I1732	Ukraine	U4b	I-L701	I2a1b1a2
I1717	5900	Unakozovskaya	Russia	J1d6	n/a (female)	n/a (female)
I8438	7300	UpperVolga	Russia	U4b1a	R-P297	R1b1a1
I8411	6930	UpperVolga	Russia	U5a1d	R-P297	R1b1a1
I12498	6500	UpperVolga	Russia	U5a2b2	R-P297	R1b1a1
I8415	5928	UpperVolga	Russia	U5a2d	n/a (female)	n/a (female)
I8405	5820	UpperVolga	Russia	U5b1b1+@16192	R-Y13200	R1b1a1a

 

[Maciamo]

The oldest R1b-Z2103 (actually Z2103 > M12149 > Y467276 > Z2106) in this study comes from the Inland Caspian region (EBA Yamnaya) and dates from 5577 years before present (c. 3600 BCE). It is sample I33307. Interestingly it is even older than the R1b-M269 and R1b-L23 in this study.

There is an even older R1b-BY95094 (R-M269 > PF7562 > FGC31929 >...> BY95094) dating from 6100 BP (c. 4100 BCE). But according to Y-Full this deep clade only formed 1100 years ago. So it's either a contamination or a mistake.

We finally have R1b-L51 (the ancestor of Italic, Celtic and Germanic R1b) in the EBA Yamna. Not many samples compared to R1b-Z2103, but it confirms that R1b-L51 also originated in the Yamna culture. They were found in very different places: Kalmykia (4900 ybp), the Don region (4600 ybp), the Volga region (4600 ybp), but also in Yamna extension as far as Romania (4850 ybp) and Serbia (4710 ybp). The Serbian sample is the only one that is also positive for L52, so that may be the first clade born outside the Pontic Steppe.

Sample I3525 is reported as R1b-Y132510 (downstream of L21 and DF13!). It comes from Hungary and dates from 4600 ybp, but it's also extremely unlikely, as that clade is estimated to be only 2900 years old. EDIT: after double checking that sample is clearly indicated as contaminated.

The oldest R1a samples are two R1a* 5M459) from Golubaya Krinitsa in the Middle Don dating from 7400 years ago (Neolithic). There are only two other R1a samples, both Z93 from the MBA Fatyanovo culture further north (an offshoot of the Corded Ware culture).

There were actually far more I2a1b-L699 (descended from Neolithic Ukrainians) in the Don region during the Yamna period than R1b-L51 or R1b-L23.

Over 300 Steppe samples and there was not a single E1b1b and only one G2a (Neolithic G2a-L91 > Z42565) from Hungary. Likewise there was only one J2b2a (L283) from Moldova. This confirms that these three haplogroups were assimilated by the Steppe people after invading Old Europe.

 

[Southpaw]

Over 300 Steppe samples and there was not a single E1b1b and only one G2a (Neolithic G2a-L91 > Z42565) from Hungary. Likewise there was only one J2b2a (L283) from Moldova. This confirms that these three haplogroups were assimilated by the Steppe people after invading Old Europe.

There is one from another paper: https://www.biorxiv.org/content/10.1101/2024.04.17.589600v1.supplementary-material

From Proto-Usatovo

I12704 Mau6 Mayaky, Kurgan 7,
Burial 211
5144±20 (weighted mean10 of
5530±32 (OxA-22959)
12,
5390±30 (BETA-441235)
5, and
5545±40 (PSUAMS-7793, this
report)
3620-3030 calBCEg M T2h2 E1b1b1a1

Among the specimens in our selection, a CTAC individual from Verteba Cave and a
proto-Usatove individual from Mayaky carried E1b1b1a1b* Y chromosomal lineages. The protoUsatove designation of the Mayaky individual is based on the 14C date that predates Usatove
even after adjusting for reservoir effect5 (Section 1.2.1). The E1b1b1a1b lineage is not
associated with the Balkan-Danubian route of farming expansion from Anatolia to central
Europe in the Neolithic137. On the other hand, the E1b1b1a1b~ expansion in the
Mediterranean, which is timed to the late Mesolithic in southeast Europe 137, raises a possibility
that it coincided with the spread of agriculture out of the Fertile Crescent into southern Europe
via the Mediterranean coast beginning ca. 9000 years ago 141. Such as, Y chromosomal lineage
diversity in the Epicardial culture of the Cardial Ware complex of the Neolithic Mediterranean
shows Near Eastern influence and Cardial/Epicardial mtDNA lineage frequencies and diversity
are comparable to those from Near Eastern Pre-Pottery Neolithic B sites142. A bearer of the
E1b1b1a1b1 haplogroup belonging to the Cardial Ware archaeological complex was reported
from the Croatian Zemunica Cave in western Balkans 8. Cardial Ware complex is considered to
have influenced the Late Neolithic-Eneolithic Hamangia culture of the west-northwest Pontic
143. Hamangia, in turn, is considered to have influenced the formation of Precucuteni-Trypillia A
144. The presence of the E1b1b1a1b1 lineage in Trypillia and a proto-Usaove individual
strengthens the link between Hamangia and CTAC as well as CTAC and Usatove and connects
the genetic ancestry of CTAC and Usatove with Cardial Ware

Burial group (kurgan) 7, Burial 2, male (25-30), 4446-4340 calBCE (5536±25 BP) (weighted mean
(Ward and Wilson, 1978): (5530±32 BP, OxA-22959); (5545±40 BP, PSUAMS-7793)]
Burial 2 of burial group (kurgan) 7 (Figure S6) was archaeologically assigned to the Usatove
culture. The grave was identified as a discoloration at the transition to the sandy loam 0.65 m
below the current surface. The bottom was at a depth of 1.25 m, the walls could be traced up
to 0.6 m. The oval burial pit was aligned from SW to NE and measured 1.4 × 1.15 m, it was
slightly longer at the bottom (1.55 × 1.15 m). The backfill was compact and consisted of three
layers: up to a depth of 1.05 m there was black earth with lenses of sandy loam, underneath
there was a 0.1 m thin layer of sandy loam, in the bottom area there was black earth with many
lime inclusions. The massive skeleton of a 25–30-year-old male laid crouched on the left side,
skull facing east and turned with the face up. The arms were bent and crossed in front of the
chest. The left hand was under the skull, the right by the facial bones. Spots of raspberry-
colored ocher were found on the skull and on the joints of the thighs and lower legs, and the
remains of organic material in the form of brown mold were also found on the lower legs.
Between the upper ribs were crumbs of dark red ocher. In front of the arms were two vessels.
One of these was a beige beaker (height 46 mm, rim diameter 45 mm, body diameter 72 mm,
base diameter 48 mm) with a squat body and low neck. Four horizontal rows of fine cord
encrusted with red ocher were applied around the neck. The body was decorated with incised
lines encrusted in white, in the upper area with a zigzag pattern forming a six-pointed star, with
three parallel grooves at regular intervals at the bottom. The clay was tempered with finely
crushed shells

A. Nikitin, S. Ivanova. Radiocarbon Dating and Stable Isotope Analysis of Human Remains from the Usatovo Culture Site of Mayaki in Ukraine

The Usatove (Usatovo) culture provides a pivotal link between the farming world of southeast Europe and the pastoralist cultures of the Pontic steppe during the Eneolithic-Early Bronze Age (EN-EBA) transition. Usatove is conventionally dated to the

www.academia.edu

 

[Archetype0ne]

The oldest R1b-Z2103 (actually Z2103 > M12149 > Y467276 > Z2106) in this study comes from the Inland Caspian region (EBA Yamnaya) and dates from 5577 years before present (c. 3600 BCE). It is sample I33307. Interestingly it is even older than the R1b-M269 and R1b-L23 in this study.

There is an even older R1b-BY95094 (R-M269 > PF7562 > FGC31929 >...> BY95094) dating from 6100 BP (c. 4100 BCE). But according to Y-Full this deep clade only formed 1100 years ago. So it's either a contamination or a mistake.

We finally have R1b-L51 (the ancestor of Italic, Celtic and Germanic R1b) in the EBA Yamna. Not many samples compared to R1b-Z2103, but it confirms that R1b-L51 also originated in the Yamna culture. They were found in very different places: Kalmykia (4900 ybp), the Don region (4600 ybp), the Volga region (4600 ybp), but also in Yamna extension as far as Romania (4850 ybp) and Serbia (4710 ybp). The Serbian sample is the only one that is also positive for L52, so that may be the first clade born outside the Pontic Steppe.

Sample I3525 is reported as R1b-Y132510 (downstream of L21 and DF13!). It comes from Hungary and dates from 4600 ybp, but it's also extremely unlikely, as that clade is estimated to be only 2900 years old. EDIT: after double checking that sample is clearly indicated as contaminated.

The oldest R1a samples are two R1a* 5M459) from Golubaya Krinitsa in the Middle Don dating from 7400 years ago (Neolithic). There are only two other R1a samples, both Z93 from the MBA Fatyanovo culture further north (an offshoot of the Corded Ware culture).

There were actually far more I2a1b-L699 (descended from Neolithic Ukrainians) in the Don region during the Yamna period than R1b-L51 or R1b-L23.

Over 300 Steppe samples and there was not a single E1b1b and only one G2a (Neolithic G2a-L91 > Z42565) from Hungary. Likewise there was only one J2b2a (L283) from Moldova. This confirms that these three haplogroups were assimilated by the Steppe people after invading Old Europe.

I believe this might have something to do with the L283:

Archetype0ne

What I understood is that the CLV people brought Anatolian IE to Anatolia from the East. And the third wave containing the CLV signal that did not directly admix with EEF like the second wave but had high CLV admix bypassed the second and went on to spread the IE languages. This is what I got from the two papers and that one podcast. Although I have to admit that I have to slowly re-read the papers as I quickly skimmed them. The R1b seems to have been the overwhelming majority only in the third wave? Not sure the composition in the first wave. But from what we know historically haplogroups do not have to mirror languages 1:1, as one is a cultural phenomenon (Basques, Turks, Magyars etc), so it is not out of the realms of possibility.

Edit: The fine print in the image here is important:

Roots of Caucasus-Lower Volga people
The people of the homeland of the Proto-Indo-Anatolians (1) had ancestry from the Caucasus Neolithic (Aknashen) and Lower Volga Eneolithic (Berezhnovka)
Caucasus hunter-gatherer ancestry was common to both groups. In the Lower Volga Eneolithic there was also Eastern hunter-gatherer (Lebyazhinka) and Central Asian-Siberian (Tutkaul) ancestry. In the Caucasus Neolithic there was also ancestry from the Neolithic people of Anatolia-Mesopotamia (Çayönü).
Pre-Indo-Anatolian languages were spoken by at least some of these diverse ancestors, but living Indo-European languages trace their origins to the Yamnaya expansion ca. 3300BCE out of (3) and the earlier Indo-Anatolian expansion ca. 4400-4000BCE out of (1)

Expansions of Caucasus-Lower Volga people
Eneolithic people of the North Caucasus-Lower Volga region (1) had a mix of Caucasus Neolithic (Aknashen) and Lower Volga Eneolithic (Berezhnovka) ancestry. Migrations from (1) to the Volga (2), Dnipro-Don (3) and Caucasus-Anatolia (4) regions spread one or both of these ancestries, admixing in (2) with Eastern hunter-gatherers (Lebyazhinka), in (3) with Dnipro-Don hunter-gatherers, and in (4) with Caucasus-Mesopotamian (Masis Blur, Çayönü) populations.
The Yamnaya were formed ca. 4000BCE in the Serednii Stiih cultural context of the Dnipro-Don area (3) before expanding ca. 3300BCE across the steppe. Anatolian speakers of Central Anatolia (4) living. ca. 2750-1500BCE shared with the Yamnaya ancestry from area (1)
The Proto-Indo-Anatolian homeland was thus Ο probably in the North Caucascus-Lower Volga Tutkaul area (1 and environs). Migrations to the west (3) and south (4) ca. 4400-4000BCE split PIA into the Proto-Indo-European languages (spoken by the Yamnaya) and Proto-Anatolian languages (spoken in Bronze Age Central Anatolia)

Armenia_Aknashen_N Armenia I1 J-PF5174 J2a1a1b1a1a
Russia_LowerVolga_EBA_Yamnaya Russia n/a (<2x) n/a (sex unknown) n/a (sex unknown)
^but maybe Volga and Upper Volga are representative:
I10363 5165 Russia_Volga_EBA_Yamnaya Russia .. n/a (female) n/a (female)
I11840 4848 Russia_Volga_EBA_Yamnaya Russia U4c1 R-Z2108 R1b1a1b1b3
I20158 4842 Russia_Volga_EBA_Yamnaya Russia n/a (<2x) R-M269 R1b1a1b
I10362 4701 Russia_Volga_EBA_Yamnaya Russia .. n/a (female) n/a (female)
I11838 4602 Russia_Volga_EBA_Yamnaya Russia U5a1a1 R-L51 R1b1a1b1a
I8438 7300 UpperVolga Russia U4b1a R-P297 R1b1a1
I8411 6930 UpperVolga Russia U5a1d R-P297 R1b1a1
I12498 6500 UpperVolga Russia U5a2b2 R-P297 R1b1a1
I8415 5928 UpperVolga Russia U5a2d n/a (female) n/a (female)
I8405 5820 UpperVolga Russia U5b1b1+@16192 R-Y13200 R1b1a1a

Extended Data Figure 1: The origin of Central Anatolian Bronze Age. (a) Fitting models include Mesopotamian (Çayönü) and steppe ancestry. (b) Models with western sources from Southeastern Europe fail except those with Mayaky or Boyanovo EBA sources both of which are Yamnaya-derived. (c) The steppe (BPgroup)+Çayönü model fails all Chalcolithic/Bronze Anatolians except Central Anatolian Bronze Age. (d) Steppe (BPgroup) ancestry observed in all individuals of the Central Anatolian Bronze Age (±3s.e. shown). (e) BPgroup-related ancestry admixed with different substrata: Aknashen-related in the North Caucasus Maikop, Masis Blur- related in Chalcolithic Armenia, and Mesopotamian-related (Çayönü) in the ancestors of the Central Asian Bronze Age, following the route (f) from the North Caucasus to Anatolia.

I4429 5300 Maikop_I4429 Russia T2a1b1a1 J-PF5119 J2a1a1a2
I3930 7535 Armenia_MasisBlur_N Armenia K1a12a1a J-Y13341 J2a1a1a2a
Mesopotamian-related (Çayönü) ??

 

[Moja]

The only difference between this study and "The genetic history of the Southern Arc" is that it says the homeland of Indo-Anatolians was a little further north in the Caucasus.

 

[MOESAN]

Thanks Maciamo.

 

[Olympus Mons]

Will there be acknowledge that I was right all along?!

So much blablabla for so many years… and in the end, it was the Shulaveri-Shomu all along, weren’t they?
And I have described the route, when and how - It took harvard, Reich and Anthony 8 years to get there?!?!

CLV is the Harvardian talk for Shulaveri-Shomu. Mentioning Aknashen is BS, it was not a culture, but he first wave of Shulaveri-Shomu. Georgia/Azerbaijan Shulaveri- Shomu were the culture.

Of course, I wrote a lot of things of the top of my head and probably wrong. – But even when, even when, 6-7 years ago I was stating (and being mocked for it) that the KUM6 girl was the Shulaveri -Shomu dispersal of the PIE via north Anatolia it looks like I was also correct.

It’s a matter of time till the South Caucasus becomes Shulaveri-Shomu as it should. Respect!

And it’s a matter of time until the Tel Tsaf samples in Israel also become Shulaveri or even that the Merimbe beni salama in Delta Nile were in fact derived from the same population. Let us see. At this point I just observe and smile.

 

[Archetype0ne]

I believe this might have something to do with the L283:

Interestingly we do have some supporting evidence on this:

I mean KDC001 and KBD001 found in the Steppe North of the Caucasus, in the same period(4kbp), kilometers apart.
[TABLE=collapse]
[TR]
[TD]

KDC001[/TD]
[TD]3824 [/TD]
[TD]RUS [RU-KB] [/TD]
[TD]J-L283 [/TD]
[TD]X2i3[/TD]
[TD]Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions[/TD]
[/TR]
[/TABLE]
[TABLE=collapse]
[TR]
[TD]

KBD001[/TD]
[TD]4037 [/TD]
[TD]RUS [RU-STA][/TD]
[TD]R1b1ba2[/TD]
[TD]I4a*[/TD]
[TD]Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions[/TD]
[/TR]
[/TABLE]

YFull | Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions

L283 in light green.

The ancient KDC001 and the modern Armenian samples sitting at ~J-L283* are in a different situation, as they don't have any European matches in the last ~5500 years and are not deep within European branches, therefore for these two it's more likely they are "native" to the wider Transcaucasian region or nearby..

Furthermore we know that NEO281 and KK1 not only lived in the same cave roughly 10kbp in Kotias Klde, but they also shared the same maternal haplogroup, and ancestry:

It looks like both NEO281 and KK1 can be modeled as approximately 80% Progress CHG. Exhibit A:

At the same time, I’ve seen some people produce models that show anywhere from 15% to 30% Meshoko ancestry contained within Progress. Even Davidski acknowledges now that this is the case over at his Eurogenes blog. That‘s why I think the Neolithic similarity map that was made by Ph2ter at GenArchivist is so interesting. It shows NEO281, a Mesolithic J2b2 sample, is most similar to the Darkveti Meshoko culture and pre Maykop people found centered there in Sochi and Adler, Russia area. These locations connect with Nal’chik to the north where the KDC001 (Nal’chik) J2b L283 sample was found. Exhibit B:

But NEO281 is actually from Kotias Klde near Imereti, Georgia. This is southeast of Sochi. So it looks to me like some J2b2 lineages moved northwest from the Imereti area up the east coast of the Black Sea via Meshoko (agro pastoralists ) and either through the mountain gorges or around the western most tip of the Caucasus into Ciscaucasia and the steppe. Even the shaded area in Ph2ter’s map wraps around the northern portion of the Black Sea extending as far west as the mouth of the Danube, but not nearly to the same extent along the southern Black Sea coast.

Finally, here is Nick Patterson’s comment on CopperAxe’s blog Musaeum Scythia from nearly 2 years ago that I think is interesting in light of the recent NEO281 sample that shows somewhat elevated Iran N ancestry relative to KK1. Exhibit C:

Thanks to Polska and Trojet for doing most of the hard lifting as far as historical research and analysis over the past few years when it comes to what is in these posts.

 

[Olympus Mons]

 

[Olympus Mons]

 

[Olympus Mons]

Just because it’s helpful to just state it again and again! - PIE was the language of the Shulaveri-Shomu.

From honey to plough to horses, they had everything. Actually, first culture that diverged a river and invented wine!
It does not matter if they call it South Caucasus, or Armenia or northwestern Iran… golly! There is no other culture in that region for 2000 years but the Shulaveri. Plenty of time to spill into the north Caucasus.



Wiki was updated on them, and it is a bit more informative. Not much but a bit. -https://en.wikipedia.org/wiki/Shulaveri–Shomu_culture

 

[Maciamo]

There is one from another paper: https://www.biorxiv.org/content/10.1101/2024.04.17.589600v1.supplementary-material

From Proto-Usatovo

I12704 Mau6 Mayaky, Kurgan 7,
Burial 211
5144±20 (weighted mean10 of
5530±32 (OxA-22959)
12,
5390±30 (BETA-441235)
5, and
5545±40 (PSUAMS-7793, this
report)
3620-3030 calBCEg M T2h2 E1b1b1a1

Yes, but the Usatovo culture is already a mix of Steppe people with the Cucuteni–Trypillia culture.

 

[Maciamo]

Will there be acknowledge that I was right all along?!

So much blablabla for so many years… and in the end, it was the Shulaveri-Shomu all along, weren’t they?
And I have described the route, when and how - It took harvard, Reich and Anthony 8 years to get there?!?!

CLV is the Harvardian talk for Shulaveri-Shomu. Mentioning Aknashen is BS, it was not a culture, but he first wave of Shulaveri-Shomu. Georgia/Azerbaijan Shulaveri- Shomu were the culture.

Of course, I wrote a lot of things of the top of my head and probably wrong. – But even when, even when, 6-7 years ago I was stating (and being mocked for it) that the KUM6 girl was the Shulaveri -Shomu dispersal of the PIE via north Anatolia it looks like I was also correct.

It’s a matter of time till the South Caucasus becomes Shulaveri-Shomu as it should. Respect!

And it’s a matter of time until the Tel Tsaf samples in Israel also become Shulaveri or even that the Merimbe beni salama in Delta Nile were in fact derived from the same population. Let us see. At this point I just observe and smile.

Do we have Y-DNA from the Shulaveri-Shomu culture?

I always thought that R1b-M269 came from the northern Middle East via the Caucasus but I thought it more likely through the eastern side of the Caucasus how long do Caspian Sea (mainly because it's easier to cross there). This remains to be confirmed by ancient DNA

 

[Southpaw]

Yes, but the Usatovo culture is already a mix of Steppe people with the Cucuteni–Trypillia culture.

In that context yeah, you are right.

 

[mount123]

Over 300 Steppe samples and there was not a single E1b1b and only one G2a (Neolithic G2a-L91 > Z42565) from Hungary. Likewise there was only one J2b2a (L283) from Moldova. This confirms that these three haplogroups were assimilated by the Steppe people after invading Old Europe.

J2b-L283 is a CHG-related steppe lineage. The sample in question is part of what the authors label Core Yamnaya. There is CHG-related auDNA in Eastern Europe since the Mesolithic. During the Chalcolithic and after it is one of the key components of the southern dry steppe. J2b-L283 obviously went to the Western Balkans through the southern dry steppe route via the Danube and its tributaries. Comparing it to ANF-related farmer lineages and hybrid-EEF cultures is erroneous.

G2a-L91 is one of the core ANF Lineages, the undefined E1b1b is ultimately just another Mesolithic Ancestral Northafrican paternal branch that got absorbed by the expanding Farmers en route to Europe.

 

[Maciamo]

J2b-L283 is a CHG-related steppe lineage. The sample in question is part of what the authors label Core Yamnaya. There is CHG-related auDNA in Eastern Europe since the Mesolithic. During the Chalcolithic and after it is one of the key components of the southern dry steppe. J2b-L283 obviously went to the Western Balkans through the southern dry steppe route via the Danube and its tributaries. Comparing it to ANF-related farmer lineages and hybrid-EEF cultures is erroneous.

That remains to be seen. This J2b-L283 sample dates from the late Yamna period (2900-2500 BCE) in Moldova (former Cucuteni-Trypillian area) and the authors say that it could also belong to the Chalcolithic Cernavodă culture. Unless I missed something from a previous study I don't think that any J2b-L283 has been found in the Pontic-Caspian Steppe itself or the North Caucasus during the Yamna and Maykop periods. Maykop had J2a, but not J2b.