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West Liao River (Bronze Age)

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Geographic location and dates of ancient individuals in Northern East Asia.
Proto-Macro-Koreanic arrived after Proto-Japanic from Liaodong and the Changbaishan region with the introduction of bronze daggers around 300 BC[1]

The Bronze Age West Liao River farmers (WLR_BA) display long-term genetic continuity with modern Koreans. Modern Koreans can be modelled to be derived primarily from Bronze Age farmers from the West Liao River.[2] West Liao River farmers of the Bronze Age themself can be modelled to be derived from the combination of two Ancient Northern East Asian lineages, namely "Neolithic Yellow River farmers" and Ancient Northeast Asians (Amur hunter-gatherers) during the Neolithic period. The spread of Proto-Koreanic can be linked to the expansion of Bronze Age West Liao River farmers. It is also suggested that this type of ancestry was introduced into the Japanese gene pool by early Koreanic-speakers, during the Kofun period.[3] WLR_BA ancestry is also associated with the Upper Xiajiadian culture, which in turn can be used as source proxy for Bronze Age and modern Koreans.[4][5]

Archaeologic evidence point to a connection between the pottery-making style of the Late Neolithic to Bronze Age cultures in the West Liao River basin and the Korean peninsula.[6]

Yayoi-Mumun

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The Yayoi people are generally associated with the Proto-Japonic-speakers and the introduction of Japonic languages into Japan during the Yayoi period from the southern Korean peninsula. They are suggested to have been closely related to the pre-Koreanic Mumun pottery period populations of the southern Korean penisular, which are linked to the presence of Peninsular Japonic.[7][8] Genetic analyses on ancient remains from southern Korea revealed elevated Jōmon ancestry at c. 37%, while Yayoi remains in Japan were found carry nearly equal amounts of Jōmon ancestry (35–60%) and Ancient Northeast Asian-like ancestry (40–65%). These results suggest the presence of a Jōmon-like population on the Korean peninsula and their significant contribution to the formation of early Japonic-speakers. As such, the "agricultural transition in prehistoric Japan involved the process of assimilation, rather than replacement, with almost equal genetic contributions from the indigenous Jomon" and mainland Asian migrants of the Mumun/Yayoi period.[9][10][11][12]

Subsequent migration waves into Japan during the Kofun period, associated with the expansion of Proto-Koreanic-speakers and Han Chinese, and the introduction of Chinese characters, saw a decline of Jōmon ancestry among modern Japanese populations to 13%–15%. A similar decline of Jōmon-like ancestry among ancient southern Korean specimens from c. 37% to nearly 0% among modern Koreans may be linked to the replacement of Peninsular Japonic speakers by early Koreanic-speakers during the Three Kingdoms period of Korea.[9][10]

Diverse archaeogenetic topics

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Cisbaikal_LNBA

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Possible Cisbaikal_LNBA affinity for Eastern Saka groups (Yeniseian layer?):[13] (supplementary)

The two ancient outliers from the Late Bronze Age Minusinsk Basin from the period of the Karasuk culture that ADMIXTURE and F4-statistics suggest have high levels of ancestry from Cisbaikal_LNBA also require such ancestry in qpAdm; interestingly, the population that succeeds the Karasuk and Lugavskaya cultures in the region, the Tagar culture (Russia_Tagar.SG), also requires ancestry from Cisbaikal_LNBA in qpAdm for all passing models. In addition, Mongolic-speaking Kalmyks and Kazakhstan_CentralKazakhSteppe_Saka also require such ancestry for passing models in this qpAdm setup, but no other populations from their ethnolinguistic or cultural categories behave similarly. All qpAdm models in this section are listed in SI Data 6, Table 4.

Fig. S80 (qpadm; supplementary); Glazkovo; Baikal EBA.

Yumin_N

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The Yumin archaeological site is located in Huade County, Ulanqab city, Inner Mongolia Autonomous Region of China. Yumin culture is the earliest Neolithic culture found in Inner Mongolia thus far. Radiocarbon analysis of charcoal samples associated with the remains of a house were dated to ~8,400 cal BP (62). We sequenced a single individual (M1) from this site, identified to be female, and she was directly radiocarbon dated to 8,415-8,335 cal BP.

Yumin: 8,500-year-old newly sampled individual from Inner Mongolia belonging to the inland nEastAsia_EN group. = distinct ANEA branch; not identical to Amur_N. Cite:[14] supplementary information

38-40% Tianyuan ancestry for Yana in supplementary models, 32% in main article (Fig. 2).

Xiongnu/Turkic/Uyghur period

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Lee & Kuang:[15]

"The geneticists who analysed the dna of the Xiongnu specimens from the Egyin Gol necropolis and that of modern Mongolians suggest that ‘the impact of the succession of Turkic and Mongolian confederations on the territory of the current Mongolia was a cultural or linguistic process rather than a migratory and/or genetic one’ (Keyser-Tracqui et al. 2006: 279). Similarly, a comparative study of the autosomal dna of the Mongols and the Tsaatan, a Turkic people residing in northern Mongolia, also concludes that the two, along with the Sakhas, form the same cluster and are genetically distinct from other world populations (Brissenden et al. 2015: 82). Finally, an extensive study of the genetic legacy of the Turkic nomads across Eurasia based on autosomal dna analysis reveals that the source populations for the Turkic nomads who spread ‘Asian genes’ to non-Turkic peoples were (the ancestors of modern-day) Tuvinians, Mongols and Buryats, despite the fact that the latter two are Mongolic (Yunusbayev et al. 2015).81 In sum, one should note that the early eastern Turkic peoples were in all likelihood genetically closer to their neighbouring Mongolic peoples than to various later Turkic peoles of central and western Eurasia.

... The analysis of genetic survey data on the Turkic peoples also allows us to speculate on the Turkic Urheimat. We suggest that it was a geographical region where the carriers of haplogroups C2, N, Q and R1a1 could intermix, since these haplogroups are carried by various past and modern-day Turkic peoples in eastern Inner Asia and the Xiongnu. It has been suggested that the early Turkic peoples probably had contact with Indo-European, Uralic, Yeniseian, and Mongolic groups in their formative period (Golden 2006: 139). As non-linguists, we are unqualified to discuss the origin of the Turkic languages. However, drawing on the findings of dna studies, we are inclined to think that certain similarities that exist between the Turkic languages and the Mongolic, Tungusic and Uralic languages are at least partly associated with haplogroups C2 and N, among others. More specifically, we conjecture that the Turkic languages came into existence as a result of the fusion of Uralic groups (characterized by a high frequency of haplogroup N subclades) and Proto-Mongolic groups (characterized by a high frequency of haplogroup C2) who also merged with other linguistic groups, including Yeniseian speakers (characterized by a high frequency of haplogroup Q like the Kets) and Indo-European speakers (characterized by a high frequency of haplogroups R1a1).

... Finally, we suggest that the Turkicisation of central and western Eurasia was the product of multiple processes of language diffusion85 that involved not only originally Turkic-speaking groups, but also Turkicised (Indo-European) groups. That is, the earliest Turkic groups first Turkicised some non-Turkic groups residing in Mongolia and beyond. Then both Turkic and ‘Turkicised’ groups Turkicised non-Turkic tribes (who were mostly carriers of haplogroups R1a1) residing in the Kazakh steppes and beyond. Through multiple processes, including the Mongol conquest, the members of the extended Turkic entity spread the Turkic languages across Eurasia."

Ancient Siberian mythology - Deer Goddess

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Deer images, carvings, paintings, and monolithic stelae of South Siberia and northern Central Asia.[16]

Indo-European

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Lazaridis et al. 2024:[17]

"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." CLV="Caucasus-Lower Volga"

Yamnaya culture = Proto-Indo-European speakers

"The Yamnaya culture stands as the unifying factor of all attested Indo-European languages. Yet, the homogeneity of the Yamnaya patrilineal community was formed out of the admixture of diverse ancestors, via proximal ancestors from the Dniproand CLVclines (Fig. 2e). Yamnaya and Anatolians share ancestry from the CLV Cline (Fig. 2e,f), and thus, if the earliest IA language speakers shared any genetic ancestry at all—the possibility of an early transfer of language without admixture must not be discounted—then the CLV Cline is where this ancestry must have come from."

CLV cline people = Indo-Anatolian; Pre-IA unknown from a genetic viewpoint:

"Genetics has little to say whether within this cline the IA languages were first spoken in the Caucasus end of the cline and spread into the steppe along with the spread of Caucasus ancestry, or vice versa, or even if a linguistic unity uncoupled with ancestry existed within the CLV continuum. DNA has traced back the ancestors of both Anatolian and IE speakers to the part of the CLV Cline that was north of the Caucasus mountains, bringing them into proximity with each other and uncovering their common CLV ancestry. However, it cannot adjudicate, on its own, who among the proximate and diverse distal ancestors of the CLV people were Pre-IA speaking."

"Aknashen" (Caucasus Neolithic) may be a relevant candidate for Pre-IA based on the presence of some cereal terminology in IA and IE, and not contradicted by genetic data on the CLV and Yamnaya. Anatolian may have arrived via an "eastern route" as opposed to the Balkan route from the CLV cline area into Anatolia (Central Anatolian Early Bronze Age people).

Iranian hunter-gatherers

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Not to be confused with Indo-Iranians or Iranian peoples.

The term Iranian hunter-gatherers or Neolithic Iranian, sometimes also "East Meta", is used to referr to a population genomics lineage representing the Mesolithic to early Neolithic population of the Iranian plateau, and to some extent regions of South-Central Asia and the Caucasus.

The Ancient Iranian lineage is represented by Mesolithic hunter-gatherers and Neolithic herders and early farmers in Iran, such as remains excavated from the Hotu and Kamarband Caves and Ganj Dareh. A deeply diverged branch of Ancient Iranians (c. 12kya) represented by remains from Shahr-i-Sokhta, formed one of the dominant ancestry components of the Indus Valley Civilisation, in tandem with an Ancient East Eurasian lineage (specifically South Asian hunter-gatherers/AASI) indigenous to South Asia. The Ancient Iranians also contributed significantly to the formation of the Central Asian gene pool, primarily via the Bactria–Margiana Archaeological Complex. They displayed close genetic affinities to the Caucasus hunter-gatherers, who derived from a similar source population as Iranian hunter-gatherers, but were distinct from preceeding Paleolithic Caucasus populations, which were closer to Anatolian hunter-gatherers.

Origins

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A possible admixture graph of deep Eurasian lineages in context of modern West Eurasians[18]
Relationship and legacy of the West and East Eurasian Core populations; WHG, Iran_N, and Levantines occupy the area below the bisector, compatible with an admixture between EEC and WEC, or below the blue axis, further complicated by the presence of Basal Eurasian or African components in these populations.
Principal Components Analysis of Ancient West Eurasians: Eigenvectors were inferred using present-day populations (gray points) and the ancient samples (colored shapes) were projected onto the plot.

While the exact origin of the Mesolithic and Neolithic Iranian hunter-gatherers and later farmers remains unclear, they are often described as having formed as combination of two deep lineages, specifically a lineage represented by Basal Eurasians and a lineage closer to Ancient North Eurasians and or Eastern European Hunter-Gatherers (EHG). In this scenario, the Mesolithic/Neolithic Iranian lineage derives significant amounts of their ancestry from Basal Eurasians (ranging from 38–48%), with their remainder ancestry being closer to Ancient North Eurasians (ANE). The related CHG displayed a higher ANE-like and Upper Paleolithic Caucasus component than the Neolithic Iranians do, suggesting contact with Eastern Hunter-Gatherers (EHG) to their North and Anatolian groups to their West. The geographically adjacent Natufians from the Levant were found to derive their ancestry primarily from the same Basal Eurasian lineage (c. 44%), but their remainder ancestry from a population closer to Western European Hunter-Gatherers (WHG).[19][20][21]

Vallini et al. 2024 presented a revised model, suggesting that Ancient Iranians (Iranian hunter-gatherers) formed primarily from a deep Ancient West Eurasian lineage ('WEC2', c. 72%), and from varying degrees of Ancient East Eurasian (c. 10%) and Basal Eurasian (c. 18%) admixture. The Ancient West Eurasian component associated with Iranian hunter-gatherers ('WEC2') is inferred to have diverged from the West Eurasian Core lineage (represented by Kostenki-14; 'WEC'), with the WEC2 component staying in the region of the Iranian Plateau, while the proper WEC component expanded into Europe and contributed to the formation of later Western Hunter-Gatherers and partially to Ancient North Eurasians.[22][a]

While Ancient Iranians fall into the wider 'West Eurasian' cluster, and display close genetic affinities to the Mesolithic Caucasus hunter-gatherers, they are only distantly related to the geographical close Anatolian or Levantine lineages, taking up an "extreme position" within a PCA of ancient and modern West Eurasian populations.[23][24][25][26]

Human Y-chromosome DNA haplogroups found among Neolithic Iranian specimens include haplogroup R2a, haplogroup CT (unknown subclade), haplogroup G2a, and haplogroup J. The oldest sample of haplogroup R2a to date was observed in one of the remains from Ganj Dareh in western Iran.[27][28][29]

Contributions to other populations

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West Asia

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The later Chalcolithic Iranians are modeled to have formed from a merger of Neolithic Iranians and a Levant and or Anatolian source population, and additional Caucasus hunter-gatherer-like geneflow.[30][31] During the Late Neolithic/Early Chalcolithic period they formed a cline stretching from Western Anatolia along the lowlands of the Southern Caucasus to the Zagros mountains, reaching as far as to Southern Central Asia, as well as southwards to the Southern Levant. This cline was primarily characterized by expansive Anatolian-like ancestry and secondarily by the spread of Neolithic Iranian and Levantine-like ancestries.[32]

A Neolithic Iranian-like contribution is needed in models for modern Middle Eastern and certain Eastern African populations. This geneflow may have happened primarily via a population from the Levant or Mesopotamia.[33]

South Asia

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A divergent (>12kya) lineage (Eastern Iranian hunter-gatherers), sharing a recent common ancestor with Neolithic Iranians, but diverging from them prior to the development of agriculture, forms one of the two main ancestry components of the Indus Valley Civilisation. The remaining ancestry is made up by a local South Asian hunter-gatherer population associated with the 'East Eurasian Core' lineage. The spread of Ancient Iranian-like ancestry may be related to the dispersal of early Dravidian languages, althought an indigenous origin and association with the 'ASI' component ("Ancestral South Indian") has been proposed as well.[34][35][28][36][37]

Central Asia

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Neolithic Iranians, in tandem with Anatolian Farmers, also contributed to the formation of the Bactria–Margiana Archaeological Complex, which subsequently contributed to other Central Asian populations, and possibly later Tarim mummies from Alwighul (700–1 BCE) and Krorän (200 CE).[38][39][40]

Europe

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Neolithic Iranians, in contrast to the related Caucasus hunter-gatherers, did only made little contributions to the European gene pool.[41] Neolithic Iranians instead represent a better source of geneflow among most West Asian populations when compared against Caucasus hunter-gatherers, while the contrary is true for European populations.[42]

Relevant:[43]

Ancient West Eurasian

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Ancient West Eurasian

The term Ancient West Eurasian, alternatively also known as West Eurasian or Western Eurasian, is used in population genomics to describe the genetic ancestry and phylogenetic relationship of diverse populations primarily living in the western and northern parts of Eurasia as well as parts of Northern and Northeastern Africa, deriving large amounts of their ancestry from the "West Eurasian Core" of human genetic diversity, and which can be associated with the Upper Paleolithic (UP) wave outgoing from Paleolithic Western Asia and Europe (eg. Kostenki-14-like WEC and WEC2), following the earlier Initial Upper Paleolithic (IUP) wave associated with the "East Eurasian Core" populatios (EEC), and ultimately the Out of Africa migration (>60kya).

Repetitive expansions into Eurasia from a population Hub OoA. (A) Zlatý Kůň can be described as a putative early expansion from the population formed after the major expansion OoA and hybridization with Neanderthals, and could be linked with non-Mousterian and non-IUP cultures found in Europe 48–45 ka or with IUP. (B) Representative samples dated between 45 and 40 ka across Eurasia can be ascribed to a population movement with uniform genetic features and material culture consistent with an IUP affiliation and which can also explain Oase1 after allowing for additional Neanderthal contributions; modern Papuans may be genetically seen as an extreme extension of this movement. (C) Following local genetic differentiation, a subsequent population expansion could explain the genetic components found in ancient samples <38 ka which contain it in unadmixed form (Kostenki14, Sunghir) or admixed with preexisting IUP components (Goyet Q116-1, Yana1, Mal’ta).
Inferred phylogenetic structure of West Eurasian lineages, taking admixture events into account.

Modern humans of the Upper Paleolithic wave (UP) associated with the "West Eurasian Core", are suggested to have expanded from a population hub located in the Iranian Plateau (c. 38kya) after an earlier "Initial Upper Paleolithic" wave. This UP wave is are linked to the "West Eurasian" ancestry represented by the Kostenki-14 specimen, and broadly ancestral to historical and modern populations in the Middle East/Western Asia, Northern and Northeastern Africa, Europe, and partially Siberia, Central Asia, and Southern Asia. Unadmixed Early West Eurasians are currently represented by several Upper Paleolithic European remains such as Kostenki-14 and Sungir. Other early Western Eurasian lineages in Europe and elsewhere displayed varying degrees of contact/admixture with preceeding IUP Ancient East Eurasian and or Basal Eurasian lineages. The expansion of early West Eurasian ancestry can be associated with Upper Paleolithic material culture distinct from previous IUP material culture.

Ancient West Eurasians can be divided into two deep early branches, specifically 'WEC' (represented by Ice Age Europeans) and 'WEC2' (making up a significant amount of ancestry of ancient Iranian hunter-gatherers. In tandem with varying amounts of East and Basal Eurasian components, these two branches gave rise to historical and modern West Eurasian lineages/populations.

Primary sources:[18][44]

Secondary sources and reviews:[45][46]

Vallini et al. 2024:

"West Eurasians, North Western South Asians, and Levantines occupy the area below the bisector, compatible with an admixture between EEC and WEC, or below the blue axis, further complicated by the presence of Basal Eurasian or African components in these populations."

"We simulated two different West Eurasian populations: WEC and WEC2, with WEC2 staying in the Hub longer than WEC (and Kostenki14), and hence closer to it from a genetic point of view. We then have each of these populations acting as a source for admixture events with Basal Eurasians (BEA) and East Eurasians in different proportions (Supplementary Data 9)."

Other

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Pre-prints:[47][17]

May be relevant:[48][49]

Secondary paper - peopling of Oceania:[50] - Glazkovo culture (Neolithic to EBA Baikal):[51]

Papuan archaic introgression:[52]

Usefull: [https://haplotree.info/maps/ancient_dna/samples.php] & [53]

IUP sites:[54]

Hub OOA:[55]

Notes

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  1. ^ Our results showed that the genetic component closest to the Hub population is represented in ancient and modern populations in the Persian Plateau. Such a component, after mixing with Basal and East Eurasian ancestries, resurfaced in the palaeogenetic record, previously referred to as the Iranian Neolithic, the Iranian Hunter Gatherer’ or the East Meta49. --> supplementary data 9 (WEA-EEA-BEA B)

Reference

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  2. ^ Sun, Na; Tao, Le; Wang, Rui; Zhu, Kongyang; Hai, Xiangjun; Wang, Chuan-Chao (2023-01-02). "The genetic structure and admixture of Manchus and Koreans in northeast China". Annals of Human Biology. 50 (1): 161–171. doi:10.1080/03014460.2023.2182912. ISSN 0301-4460. Koreans can also be modelled as deriving ancestry from a single source related to WLR_BA, consisting of the transmission route of farming from the northeast to the Korean Peninsula and even the Japanese islands (Kwak et al. 2017; Kim and Park 2020).
  3. ^ Wang, Rui; Wang, Chuan-Chao (2022-08-08). "Human genetics: The dual origin of Three Kingdoms period Koreans". Current Biology. 32 (15): R844–R847. doi:10.1016/j.cub.2022.06.044. ISSN 0960-9822. The northern East Asian ancestry was suggested to be related to the Neolithic West Liao River farmers in northeast China, who were an admixture of ANA and NYR ancestry3. The finding indicated that West Liao River-related farmers might have spread the proto-Korean language as their ancestry was found to be predominant in extant Koreans. Proto-Korean groups, in turn, introduced West Liao River-like ancestry into the gene pool of present-day Japan5. {{cite journal}}: no-break space character in |title= at position 35 (help)
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  18. ^ a b Allentoft, Morten E.; Sikora, Martin; Refoyo-Martínez, Alba; Irving-Pease, Evan K.; Fischer, Anders; Barrie, William; Ingason, Andrés; Stenderup, Jesper; Sjögren, Karl-Göran; Pearson, Alice; Sousa da Mota, Bárbara; Schulz Paulsson, Bettina; Halgren, Alma; Macleod, Ruairidh; Jørkov, Marie Louise Schjellerup (2024-01). "Population genomics of post-glacial western Eurasia". Nature. 625 (7994): 301–311. doi:10.1038/s41586-023-06865-0. ISSN 1476-4687. {{cite journal}}: Check date values in: |date= (help)
  19. ^ Jones, Eppie R.; Gonzalez-Fortes, Gloria; Connell, Sarah; Siska, Veronika; Eriksson, Anders; Martiniano, Rui; McLaughlin, Russell L.; Gallego Llorente, Marcos; Cassidy, Lara M.; Gamba, Cristina; Meshveliani, Tengiz; Bar-Yosef, Ofer; Müller, Werner; Belfer-Cohen, Anna; Matskevich, Zinovi (2015-11-16). "Upper Palaeolithic genomes reveal deep roots of modern Eurasians". Nature Communications. 6 (1): 8912. doi:10.1038/ncomms9912. ISSN 2041-1723.
  20. ^ Lazaridis, Iosif; Nadel, Dani; Rollefson, Gary; Merrett, Deborah C.; Rohland, Nadin; Mallick, Swapan; Fernandes, Daniel; Novak, Mario; Gamarra, Beatriz; Sirak, Kendra; Connell, Sarah; Stewardson, Kristin; Harney, Eadaoin; Fu, Qiaomei; Gonzalez-Fortes, Gloria (2016-08). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. doi:10.1038/nature19310. ISSN 1476-4687. Neolithic Iran and Natufians could be derived from the same Basal Eurasian population but are genetically closer to EHG and WHG respectively. We take the model of Fig. S4.9 and attempt to fit Natufians as a mixture of the same Basal Eurasian population that contributes to Iran_N and any other population of the tree. Several solutions are feasible, and we show the best one (lowest ADMIXTUREGRAPH score) in Fig. S4.10. We can add both EHG and MA1 as simple branches to the model structure of Fig. S4.10 and show the results in Fig. S4.11. An interesting aspect of this model is that it derives both Natufians and Iran_N from Basal Eurasians but Natufians have ancestry from a population related to WHG, while Iran_N has ancestry related to EHG. Natufians and Iran_N may themselves reside on clines of WHG-related/EHG-related admixture. {{cite journal}}: Check date values in: |date= (help)
  21. ^ Almarri, Mohamed A.; Haber, Marc; Lootah, Reem A.; Hallast, Pille; Al Turki, Saeed; Martin, Hilary C.; Xue, Yali; Tyler-Smith, Chris (2021-09). "The genomic history of the Middle East". Cell. 184 (18): 4612–4625.e14. doi:10.1016/j.cell.2021.07.013. ISSN 0092-8674. PMC 8445022. PMID 34352227. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
  22. ^ Vallini, Leonardo; Zampieri, Carlo; Shoaee, Mohamed Javad; Bortolini, Eugenio; Marciani, Giulia; Aneli, Serena; Pievani, Telmo; Benazzi, Stefano; Barausse, Alberto; Mezzavilla, Massimo; Petraglia, Michael D.; Pagani, Luca (2024-03-25). "The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal". Nature Communications. 15 (1): 1882. doi:10.1038/s41467-024-46161-7. ISSN 2041-1723. We simulated two different West Eurasian populations: WEC and WEC2, with WEC2 staying in the Hub longer than WEC (and Kostenki14), and hence closer to it from a genetic point of view. We then have each of these populations acting as a source for admixture events with Basal Eurasians (BEA) and East Eurasians in different proportions (Supplementary Data 9). ... Using msprime45, we performed coalescent simulations to obtain the distinct WEC source populations under different demographic scenarios46,47 (Methods, Supplementary Code 1–2, Supplementary Fig. 4). In particular, we simulated WEC populations with different allele sharing with Kostenki14 (i.e., mimicking WEC populations with different distances from the Hub population) and mixed them with the EEC and Basal Eurasians (Supplementary Data 9). We found that our approach retrieves the correct ranking along the K coordinate in the majority of cases, and with an accuracy of >0.9 in all cases where the admixed populations are at least 50% WEC, and the mixing WEC sources have at least 3 ky of differential allele sharing with Kostenki14 (Supplementary Fig. 5A). ... We found that after accounting for East and Basal Eurasian confounders, the populations that harbour the WEC component closer to the Hub population (grayscale gradient of population points in Fig. 2A, Supplementary Data 11) are the ones whose West Eurasian ancestry is related to the hunter gatherers and early farmers from Iran48. ... Along the blue axis of genetic similarity to Kostenki14, these populations come before modern and ancient groups from the Levant and, in turn, before groups from Europe and other areas associated with the Anatolian Neolithic expansion49,51,52,53. The furthermost groups along this axis are post- and pre-LGM European hunter gatherers, which is expected owing to their genetic proximity to Kostenki14.
  23. ^ Lazaridis, Iosif; et al. (16 November 2023). "Paleolithic DNA from the Caucasus reveals core of West Eurasian ancestry". doi:10.1101/423079. Supplementary: More data from the Caucasus and Iranian plateau and Siberia/Central Asia may improve our understanding of both the Dzudzuana-related ancestors of these populations (that may have differed from Dzudzuana, e.g., in the proportion of Deep ancestry), and also of the eastern influences. ... Both CHG and Iran_N can fit as a 4-way mixure with Mbuti as one source, Dzudzuana as another, and a combination of eastern non-African (ENA) and Ancient North Eurasian (ANE) ancestry. >50% of the ancestry is inferred to derive from Dzudzuana in Iran_N and >64% in CHG. Previously we had shown that CHG could be modelled as a mixture of Iran_N and European huntergatherers12. The Dzudzuana population clarifies the origin of these populations by showing that European affinity in the Caucasus decreased between Dzudzuana at ~26 kya and Satsurblia at ~13 kya as additional ENA/ANE ancestry arrived. Thus, Iran_N/CHG are seen as descendants of populations that existed in the Villabruna→Basal Eurasian cline alluded to above, but with extra Basal Eurasian ancestry (compared to Dzudzuana), and also with ENA/ANE ancestry. The extra ENA/ANE ancestry also explains the affinity between Iran/Caucasus and EHG previously proposed as part of a North/East West Eurasian interaction sphere12, which our results suggest was created by admixture of ENA/ANE ancestry on top of the Villabruna→Basal Eurasian cline. In the north, Karelia_HG traces its ancestry to a Villabruna-related source modified by ENA/ANE admixture, while CHG/Iran_N were Dzudzuana+Basal Eurasian (or, equivalently Villabruna+Basal Eurasian) derived populations also modified by ENA/ANE admixture. As seen above, populations of mixed ENA/ANE admixture (such as Russia_Baikal_EN) already existed in Siberia by the Neolithic, although with a preponderance of ENA over ANE ancestry (the opposite of what we see in the eastern periphery of West Eurasia from Eastern Europe to Iran).
  24. ^ Lazaridis, Iosif; Alpaslan-Roodenberg, Songül; Acar, Ayşe; Açıkkol, Ayşen; Agelarakis, Anagnostis; Aghikyan, Levon; Akyüz, Uğur; Andreeva, Desislava; Andrijašević, Gojko; Antonović, Dragana; Armit, Ian; Atmaca, Alper; Avetisyan, Pavel; Aytek, Ahmet İhsan; Bacvarov, Krum (2022-08-26). "Ancient DNA from Mesopotamia suggests distinct Pre-Pottery and Pottery Neolithic migrations into Anatolia". Science. 377 (6609): 982–987. doi:10.1126/science.abq0762. ISSN 0036-8075. PMC 9983685. PMID 36007054. Supplementary: We note that one of the Test populations, the Neolithic population of the Zagros from Iran(1) cannot be well-modeled with either 1 or 2 of the Sources, consistent with its extreme PCA position in the context of West Eurasian variation.{{cite journal}}: CS1 maint: PMC format (link)
  25. ^ Lazaridis, Iosif; Nadel, Dani; Rollefson, Gary; Merrett, Deborah C.; Rohland, Nadin; Mallick, Swapan; Fernandes, Daniel; Novak, Mario; Gamarra, Beatriz; Sirak, Kendra; Connell, Sarah; Stewardson, Kristin; Harney, Eadaoin; Fu, Qiaomei; Gonzalez-Fortes, Gloria (2016-08-25). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. doi:10.1038/nature19310. ISSN 0028-0836. PMC 5003663. PMID 27459054. Western Iranian first farmers cluster with the likely Mesolithic HotuIIIb individual and more remotely with hunter-gatherers from the southern Caucasus (Fig. 1b)
  26. ^ Lazaridis, Iosif; Alpaslan-Roodenberg, Songül; Acar, Ayşe; Açıkkol, Ayşen; Agelarakis, Anagnostis; Aghikyan, Levon; Akyüz, Uğur; Andreeva, Desislava; Andrijašević, Gojko; Antonović, Dragana; Armit, Ian; Atmaca, Alper; Avetisyan, Pavel; Aytek, Ahmet İhsan; Bacvarov, Krum (2022-08-26). "Ancient DNA from Mesopotamia suggests distinct Pre-Pottery and Pottery Neolithic migrations into Anatolia". Science. 377 (6609): 982–987. doi:10.1126/science.abq0762. ISSN 0036-8075. PMC 9983685. PMID 36007054. Supplementary: Within the inland cluster, individuals that are more geographically distant from the Mediterranean, such as those from the South Caucasus [Caucasus hunter-gatherers from Georgia (10) and Ganj Dareh from Central Zagros], are also genetically more distant as compared with the geographically and genetically intermediate individuals from Mesopotamia and Armenia/Azerbaijan.{{cite journal}}: CS1 maint: PMC format (link)
  27. ^ Lazaridis, Iosif; Nadel, Dani; Rollefson, Gary; Merrett, Deborah C.; Rohland, Nadin; Mallick, Swapan; Fernandes, Daniel; Novak, Mario; Gamarra, Beatriz; Sirak, Kendra; Connell, Sarah; Stewardson, Kristin; Harney, Eadaoin; Fu, Qiaomei; Gonzalez-Fortes, Gloria (2016-08-25). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. doi:10.1038/nature19310. ISSN 0028-0836. PMC 5003663. PMID 27459054. Western Iranian first farmers cluster with the likely Mesolithic HotuIIIb individual and more remotely with hunter-gatherers from the southern Caucasus (Fig. 1b)
  28. ^ a b Narasimhan, Vagheesh M.; Patterson, Nick; Moorjani, Priya; Rohland, Nadin; Bernardos, Rebecca; Mallick, Swapan; Lazaridis, Iosif; Nakatsuka, Nathan; Olalde, Iñigo; Lipson, Mark; Kim, Alexander M.; Olivieri, Luca M.; Coppa, Alfredo; Vidale, Massimo; Mallory, James (2019-09-06). "The formation of human populations in South and Central Asia". Science. 365 (6457). doi:10.1126/science.aat7487. ISSN 0036-8075. PMC 6822619. PMID 31488661.{{cite journal}}: CS1 maint: PMC format (link)
  29. ^ "haplotree.info - ancientdna.info. Map based on All Ancient DNA v. 2.07.26". haplotree.info. Retrieved 2024-02-23.
  30. ^ Lazaridis, Iosif; Nadel, Dani; Rollefson, Gary; Merrett, Deborah C.; Rohland, Nadin; Mallick, Swapan; Fernandes, Daniel; Novak, Mario; Gamarra, Beatriz; Sirak, Kendra; Connell, Sarah; Stewardson, Kristin; Harney, Eadaoin; Fu, Qiaomei; Gonzalez-Fortes, Gloria (2016-08-25). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. doi:10.1038/nature19310. ISSN 0028-0836. PMC 5003663. PMID 27459054. The Chalcolithic people of western Iran can be modelled as a mixture of the Neolithic people of western Iran, the Levant, and Caucasus Hunter Gatherers (CHG), consistent with their position in the PCA (Fig. 1b).
  31. ^ Skourtanioti, Eirini; Erdal, Yilmaz S.; Frangipane, Marcella; Balossi Restelli, Francesca; Yener, K. Aslıhan; Pinnock, Frances; Matthiae, Paolo; Özbal, Rana; Schoop, Ulf-Dietrich; Guliyev, Farhad; Akhundov, Tufan; Lyonnet, Bertille; Hammer, Emily L.; Nugent, Selin E.; Burri, Marta (2020-05-28). "Genomic History of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus". Cell. 181 (5): 1158–1175.e28. doi:10.1016/j.cell.2020.04.044. ISSN 0092-8674. Iran_C itself can be modeled as a mixture of Iran_N and Barcın_N (p = 0.365; 37% ± 3% from Barcın_N)
  32. ^ Skourtanioti, Eirini; Erdal, Yilmaz S.; Frangipane, Marcella; Balossi Restelli, Francesca; Yener, K. Aslıhan; Pinnock, Frances; Matthiae, Paolo; Özbal, Rana; Schoop, Ulf-Dietrich; Guliyev, Farhad; Akhundov, Tufan; Lyonnet, Bertille; Hammer, Emily L.; Nugent, Selin E.; Burri, Marta (2020-05-28). "Genomic History of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus". Cell. 181 (5): 1158–1175.e28. doi:10.1016/j.cell.2020.04.044. ISSN 0092-8674. We describe a Late Neolithic/Early Chalcolithic (6th millennium BCE) genetic cline stretching from Western Anatolia (i.e., area around the Sea of Marmara) to the lowlands of the Southern Caucasus that was formed by an admixture process that started at the beginning of Late Neolithic (∼6500 years BCE). The eastern end of this cline extends beyond the Zagros mountains with minute proportions of Anatolian (i.e., Western Anatolian-like) ancestry reaching as far as Chalcolithic and Bronze Age Central Asia (Narasimhan et al., 2019). To the south, Anatolian ancestry is present in the Southern Levantine Neolithic populations (Lazaridis et al., 2016), and to the north, in the Chalcolithic and Bronze Age populations from the Caucasus (mainly mountainous area) (Allentoft et al., 2015, Lazaridis et al., 2016, Wang et al., 2019), most likely as a result of the Late Neolithic admixture.
  33. ^ Almarri, Mohamed A.; Haber, Marc; Lootah, Reem A.; Hallast, Pille; Al Turki, Saeed; Martin, Hilary C.; Xue, Yali; Tyler-Smith, Chris (2021-09-02). "The genomic history of the Middle East". Cell. 184 (18): 4612–4625.e14. doi:10.1016/j.cell.2021.07.013. ISSN 0092-8674. PMC 8445022. PMID 34352227. An additional source of ancestry needed to model modern Middle Easterners is related to ancient Iranians. Our admixture tests show that this ancestry first reached the Levant and subsequently reached Arabia and East Africa. ... suggesting a potential population carrying this ancestry (possibly unsampled yet from the Levant or Mesopotamia).
  34. ^ Gallego-Llorente, M.; Connell, S.; Jones, E. R.; Merrett, D. C.; Jeon, Y.; Eriksson, A.; Siska, V.; Gamba, C.; Meiklejohn, C.; Beyer, R.; Jeon, S.; Cho, Y. S.; Hofreiter, M.; Bhak, J.; Manica, A. (2016-08-09). "The genetics of an early Neolithic pastoralist from the Zagros, Iran". Scientific Reports. 6 (1): 31326. doi:10.1038/srep31326. ISSN 2045-2322.
  35. ^ Shinde, Vasant; Narasimhan, Vagheesh M.; Rohland, Nadin; Mallick, Swapan; Mah, Matthew; Lipson, Mark; Nakatsuka, Nathan; Adamski, Nicole; Broomandkhoshbacht, Nasreen; Ferry, Matthew; Lawson, Ann Marie; Michel, Megan; Oppenheimer, Jonas; Stewardson, Kristin; Jadhav, Nilesh (2019-10-17). "An Ancient Harappan Genome Lacks Ancestry from Steppe Pastoralists or Iranian Farmers". Cell. 179 (3): 729–735.e10. doi:10.1016/j.cell.2019.08.048. ISSN 0092-8674.
  36. ^ Broushaki, Farnaz; Thomas, Mark G; Link, Vivian; López, Saioa; van Dorp, Lucy; Kirsanow, Karola; Hofmanová, Zuzana; Diekmann, Yoan; Cassidy, Lara M.; Díez-del-Molino, David; Kousathanas, Athanasios; Sell, Christian; Robson, Harry K.; Martiniano, Rui; Blöcher, Jens (2016-07-29). "Early Neolithic genomes from the eastern Fertile Crescent". Science (New York, N.Y.). 353 (6298): 499–503. doi:10.1126/science.aaf7943. ISSN 0036-8075. PMC 5113750. PMID 27417496.
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  38. ^ Gallego-Llorente, M.; Connell, S.; Jones, E. R.; Merrett, D. C.; Jeon, Y.; Eriksson, A.; Siska, V.; Gamba, C.; Meiklejohn, C.; Beyer, R.; Jeon, S.; Cho, Y. S.; Hofreiter, M.; Bhak, J.; Manica, A. (2016-08-09). "The genetics of an early Neolithic pastoralist from the Zagros, Iran". Scientific Reports. 6 (1): 31326. doi:10.1038/srep31326. ISSN 2045-2322.
  39. ^ Narasimhan, Vagheesh M.; Patterson, Nick; Moorjani, Priya; Rohland, Nadin; Bernardos, Rebecca; Mallick, Swapan; Lazaridis, Iosif; Nakatsuka, Nathan; Olalde, Iñigo; Lipson, Mark; Kim, Alexander M.; Olivieri, Luca M.; Coppa, Alfredo; Vidale, Massimo; Mallory, James (2019-09-06). "The formation of human populations in South and Central Asia". Science. 365 (6457). doi:10.1126/science.aat7487. ISSN 0036-8075. PMC 6822619. PMID 31488661.{{cite journal}}: CS1 maint: PMC format (link)
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  41. ^ Gallego-Llorente, M.; Connell, S.; Jones, E. R.; Merrett, D. C.; Jeon, Y.; Eriksson, A.; Siska, V.; Gamba, C.; Meiklejohn, C.; Beyer, R.; Jeon, S.; Cho, Y. S.; Hofreiter, M.; Bhak, J.; Manica, A. (2016-08-09). "The genetics of an early Neolithic pastoralist from the Zagros, Iran". Scientific Reports. 6 (1): 31326. doi:10.1038/srep31326. ISSN 2045-2322.
  42. ^ Skourtanioti, Eirini; Erdal, Yilmaz S.; Frangipane, Marcella; Balossi Restelli, Francesca; Yener, K. Aslıhan; Pinnock, Frances; Matthiae, Paolo; Özbal, Rana; Schoop, Ulf-Dietrich; Guliyev, Farhad; Akhundov, Tufan; Lyonnet, Bertille; Hammer, Emily L.; Nugent, Selin E.; Burri, Marta (2020-05-28). "Genomic History of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus". Cell. 181 (5): 1158–1175.e28. doi:10.1016/j.cell.2020.04.044. ISSN 0092-8674. Therefore, it seems to hold that ancient Iranian groups overall serve as a better proxy than the Caucasus groups, although higher resolution data are necessary to compare them further.
  43. ^ Martiniano, Rui; Haber, Marc; Almarri, Mohamed A.; Mattiangeli, Valeria; Kuijpers, Mirte C.M.; Chamel, Berenice; Breslin, Emily M.; Littleton, Judith; Almahari, Salman; Aloraifi, Fatima; Bradley, Daniel G.; Lombard, Pierre; Durbin, Richard (2024-02). "Ancient genomes illuminate Eastern Arabian population history and adaptation against malaria". Cell Genomics: 100507. doi:10.1016/j.xgen.2024.100507. ISSN 2666-979X. {{cite journal}}: Check date values in: |date= (help)
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