Iranian hunter-gatherers

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

The term Iranian hunter-gatherers or Neolithic Iranians, sometimes also "East Meta", is used to refer to a population genomics lineage representing the Mesolithic to early Neolithic population of the Iranian plateau, South-Central Asia and the Caucasus.[1]

The Iranian hunter-gatherer lineage is represented by Mesolithic hunter-gatherers and later Neolithic herders and early farmers in present-day Iran, such as remains excavated from the Hotu and Kamarband Caves and Ganj Dareh, as well as Wezmeh. A deeply diverged sister branch (> 12kya) best represented by remains from Shahr-i-Sokhta, formed the dominant ancestry component of the Indus Valley Civilisation in Northwestern India, which was mixed with a local East Eurasian component termed Ancient Ancestral South Indian (AASI). The Iranian hunter-gatherers also represent an important source for the formation of the Central Asian gene pool, primarily via the Bactria–Margiana Archaeological Complex. They further displayed close genetic affinities to the Caucasus hunter-gatherers, who descend primarily from a similar source population as Iranian hunter-gatherers, but were distinct from preceding Paleolithic Caucasus populations, which were closer related to Anatolian hunter-gatherers, Western hunter-gatherers and Levantine groups.

Origins

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One possible admixture graph of deep Eurasian lineages in context of modern West Eurasians.[2]
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.

Iranian-hunter gatherers are inferred to have originated from the local Upper Paleolithic population of the Persian plateau. The Persian plateau acted as population hub during the initial colonisation of Eurasia after the Out-of-Africa expansion of modern humans (between 70–46kya). The region is also close to the area which may have habored the Basal Eurasian lineage, which is supposed to have been centered in the now sunken Persian Gulf.[1][3][4][5]

While Iranian hunter-gatherers are broadly placed as part of the wider "West Eurasian" cluster, the exact origin of the Mesolithic and Neolithic Iranian hunter-gatherers and later farmers remains unclear. Mesolithic hunter-gatherer remains from the Alborz mountain range were found to show ancestry primarily related to Basal Eurasians (c. 66%), and are ancestral to later Neolithic herders and farmers.[4][6][7][8][9][5]

The later Neolithic Iranians are commonly modeled as two-way admixture between a Basal Eurasian lineage and a lineage closer to Ancient North Eurasians (ANE) or Eastern European Hunter-Gatherers (EHG). Accordingly, the Mesolithic/Neolithic Iranian lineage derives the majority of their ancestry from a Basal Eurasian-like source (ranging from 48–68%), with the remainder ancestry being closer to Ancient North Eurasians (32–52%).[10][6][11][2][5][12]

Mesolithic and Neolithic Iranians fall along a cline between Basal Eurasian and ANE/EHG-related ancestry. This stands in contrast to Neolithic Anatolian and Levant groups, who fall along a cline between Basal Eurasian and WHG-related ancestry. Natufians, despite sharing most of their ancestry with Neolithic Anatolian groups, have an additional "Ancient North African" (ANA) ancestry component.[5][6] This distinction is also evident in that Iranian hunter-gatherers and Neolithic Iranian groups are taking up an "extreme position" compared against other ancient and modern West Eurasian populations within an Eurasian-wide Principal component analysis (PCA).[13][14][6][5]

Vallini et al. (2024) argued that part of the West Eurasian component (WEC2) among Neolithic Iranians stayed closer to the population hub than compared to Upper Paleolithic Europeans. After contact events with Basal and East Eurasian ancestry, this lineage resurfaced in the palaeogenetic record as the Iranian Neolithic, the Iranian Hunter Gatherer’ or the "East Meta".[1] Neolithic Iranians can also be modeled via a three-way admixture, including an Upper Paleolithic Caucasus/Dzudzuana-like source (itself an admixture between c. 24–28% Basal Eurasian and c. 72–76% Upper Paleolithic European ancestry; and close to later Anatolian hunter-gatherers), an ANE-like source, and an additional Basal Eurasian source.[13][5]

The f4-ratio estimation for the total Basal Eurasian ancestry among Iranian hunter-gatherers and later herders or early farmers is c. 66–68% and 58–59% respectively.[6]

While initially absent from Anatolia, Mesopotamia and the Caucasus, as well as India, it reached these regions via the expansion of Mesolithic and Neolithic groups, resulting in a cline between Iran Neolithic-like and local sources.[5]

Uniparental haplogroups

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Hotu cave dwellers in Alborz mountain range were identified as Iranian hunter gatherer
Ganj Dareh herders were of Iranian hunter gatherer ancestry

The main human Y-chromosome DNA haplogroups found among Mesolithic and Neolithic Iranian-affiliated specimens include subclades of G-M201, J, L, and R2. Others included subclades of T-M184, while subclades of H are observed among 'IVC periphery' like remains. The oldest sample of haplogroup R2a to date has been found in one of the remains from Ganj Dareh in western Iran. Common Human mitochondrial DNA haplogroups found among Mesolithic and Neolithic Iranian specimens include subclades of haplogroup U, HV, X, R, H, W, T, and M.[15][7][16][5]

Contributions to other populations

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Caucasus

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Caucasus hunter-gatherers were found to be distinct from the earlier Upper Paleolithic Caucasus/Dzudzuana population, but closely related to Mesolithic and Neolithic Iranians. They are inferred to descend primarily from the same source population, which reached the Caucasus region sometimes betwee 25kya and 13kya. They can be used as interchangeable source for Holocene populations, with various names have been created to group them together, such as Iran_N/CHG, Iran/Caucasus or Zagros/Caucasus ancestry.[17][1] The CHG can be modeled as merger between an Iranian hunter-gatherer related source (c. 72%), an UP Caucasus (Dzudzuana) related source (c. 18%), and an Eastern hunter-gatherer related source (c. 10%).[18][2] An alternative model suggests c. 72% from an Iranian hunter-gatherer source, c. 21% from an EHG-like source, and c. 7% from a WHG-like source.[6]

West Asia

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The later Chalcolithic Iranians are modeled to have formed from a merger of local Neolithic Iranians and a Neolithic Levantin-like source population, as well as additional Caucasus hunter-gatherer-like geneflow. Chalcolithic Iranians can be modeled to derive c. 80% of their ancestry from a Neolithic Iranian-like group and c. 20% from a Neolithic Levantine-like group (which itself carried Neolithic Anatolian and Natufian components). Alternatively, they can be modeled as two-way admixture between c. 87% CHG-like and 13% Neolithic Levantine-like, or as three-way admixture between c. 17% Iranian hunter-gatherers, 63% CHG-like, and 20% Neolithic Levantine-like.[6][19]

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 and Iranian-like ancestries and secondarily by the spread of Levantine-like ancestry.[20] Chalcolithic Iranian groups had a wide impact on Chalcolithic Anatolians and Bronze Age Levantine groups, contributing 33% and 44% ancestry respectively.[6]

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 an admixed population from the Mesopotamia.[21]

South Asia

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An Ancient Harappan Genome

In 2019, the genome sequence of a skeleton remain from a cemetery near Rakhigarhi dating to around 2,800-2,300 BCE, suggesting that the majority of the genome was related to Iranian hunter-gatherers. The divergent (>12kya) sister lineage, sharing a recent common ancestor with Neolithic Iranians, but having diverged from them prior to the development of agriculture, best represented by remains from Shahr-i-Sokhta, forms the main ancestry component of the Indus Valley Civilisation, in tandem with variable amounts of a local East Eurasian ancestry termed Ancient Ancestral South Indian (AASI). The spread of Ancient Iranian-like ancestry, and or IVC-like ancestry, may be related to the dispersal of early Dravidian languages, although this remains uncertain, with opposing views having been presented as well, with some scholars connecting this ancestry with the spread of Indo-Aryan languages.[22][8][7][12][23][24][25][26]

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).[22][7][27]

Europe

[edit]

Neolithic Iranians, in contrast to the related Caucasus hunter-gatherers, only made minor contributions to the European gene pool.[22] 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.[28]

See also

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References

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  1. ^ a b c d 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 (25 March 2024). "The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal". Nature Communications. 15 (1): 1882. Bibcode:2024NatCo..15.1882V. doi:10.1038/s41467-024-46161-7. ISSN 2041-1723. PMC 10963722. PMID 38528002. We simulate the OOA 60 kya, with Basal Eurasians (BEA in Supplementary Fig. 4) splitting soon after (57.5 kya) and the split between EEC and WEC, with the former leaving the Hub18, 46 kya (allowing the time for them to reach Ust'Ishim and Bacho Kiro by ~45 kya). [...] 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. [...] This is a genetic ancestry commonly referred to as the Iran Neolithic30 or the East Meta49, here named Iran HG for clarity (Supplementary Data 11). The Iran HG ancestry is widespread not only in modern-day Iran but also across ancient and modern samples from the Caucasus (in particular in the Mesolithic hunter gatherers of that region) and in the northwestern part of South Asia50.
  2. ^ a b c 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 (10 January 2024). "Population genomics of post-glacial western Eurasia". Nature. 625 (7994): 301–311. Bibcode:2024Natur.625..301A. doi:10.1038/s41586-023-06865-0. ISSN 1476-4687. PMC 10781627. PMID 38200295.
  3. ^ Ferreira, Joana C; Alshamali, Farida; Montinaro, Francesco; Cavadas, Bruno; Torroni, Antonio; Pereira, Luisa; Raveane, Alessandro; Fernandes, Veronica (1 September 2021). "Projecting Ancient Ancestry in Modern-Day Arabians and Iranians: A Key Role of the Past Exposed Arabo-Persian Gulf on Human Migrations". Genome Biology and Evolution. 13 (9): evab194. doi:10.1093/gbe/evab194. ISSN 1759-6653. PMC 8435661. PMID 34480555.
  4. ^ a b Amjadi, Motahareh Ala; Özdemir, Yusuf Can; Ramezani, Maryam; Jakab, Kristóf; Megyes, Melinda; Bibak, Arezoo; Salehi, Zeinab; Hayatmehar, Zahra; Taheri, Mohammad Hossein (4 February 2025), Ancient DNA indicates 3,000 years of genetic continuity in the Northern Iranian Plateau, from the Copper Age to the Sassanid Empire, bioRxiv, doi:10.1101/2025.02.03.636298, retrieved 17 April 2025 Quote:"Mesolithic hunter-gatherer (HG) remains from the Alborz Mountains in northern Iran show ancestries primarily related to the Basal Eurasian5,12."
  5. ^ a b c d e f g h Chataigner, Christine (1 December 2024). "The South Caucasus from the Upper Palaeolithic to the Neolithic: Intersection of the genetic and archaeological data". Quaternary Science Reviews. 345: 109061. Bibcode:2024QSRv..34509061C. doi:10.1016/j.quascirev.2024.109061. ISSN 0277-3791. The populations of the Caucasus became differentiated from those of southeastern Europe through mixing with people having Basal Eurasian ancestry. This hypothetical lineage of modern humans having almost no Neanderthal ancestry would have been first established (before 50. ka cal BP) in the region of the Persian Gulf when it was not flooded (Ferreira et al., 2021) and would also have been one of the main ancestral lines (Western Asia_UP) of the populations of Iran (Allentoft et al., 2024: Fig. S3d.16). [...] It therefore appears that the ancestral population of CHG and Iran_N (the latter genome being composed of Basal Eurasian and Ancient North Eurasian sources; Broushaki et al., 2016; Allentoft et al., 2024) shows a high genetic difference at this time compared to those of Anatolia and the Levant. This may suggest that the Caucasus-Iran group and the Anatolia-Levant group remained isolated from each other during the LGM and evolved separately (Jones et al., 2015; Altınışık et al., 2022; Guarino-Vignon et al., 2023). [...] Natufians (Raqefet cave) would have resulted, like Caucasus_UP, from a mixture of West Eurasian and Basal Eurasian, but they would also have received a contribution from Ancestral North African (Lazaridis et al., 2018).
  6. ^ a b c d e f g h 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 (25 July 2016). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. Bibcode:2016Natur.536..419L. doi:10.1038/nature19310. ISSN 1476-4687. PMC 5003663. PMID 27459054. We used qpAdm7 to estimate Basal Eurasian ancestry in each Test population. We obtain the highest estimates in the earliest populations from both Iran (66±13% in the likely Mesolithic sample, 48±6% in Neolithic samples), and the Levant (44±8% in Epipaleolithic Natufians) (Fig. 2), showing that Basal Eurasian ancestry was widespread across the ancient Near East. [...] 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.
  7. ^ a b c d 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 (6 September 2019). "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.
  8. ^ a b 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 (17 October 2019). "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. PMC 6800651. PMID 31495572.
  9. ^ 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 (1 September 2024). "Population genomics of post-glacial western Eurasia". Nature. 625 (7994): 301–311. Bibcode:2024Natur.625..301A. doi:10.1038/s41586-023-06865-0. ISSN 1476-4687. PMC 10781627. PMID 38200295.
  10. ^ 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 (16 November 2015). "Upper Palaeolithic genomes reveal deep roots of modern Eurasians". Nature Communications. 6 (1): 8912. Bibcode:2015NatCo...6.8912J. doi:10.1038/ncomms9912. hdl:11392/2418166. ISSN 2041-1723. PMC 4660371. PMID 26567969.
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  14. ^ 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 (26 August 2022). "Ancient DNA from Mesopotamia suggests distinct Pre-Pottery and Pottery Neolithic migrations into Anatolia". Science. 377 (6609): 982–987. Bibcode:2022Sci...377..982L. 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. [...] 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.
  15. ^ 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 (25 August 2016). "Genomic insights into the origin of farming in the ancient Near East". Nature. 536 (7617): 419–424. Bibcode:2016Natur.536..419L. 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)
  16. ^ "haplotree.info - ancientdna.info. Map based on All Ancient DNA v. 2.07.26". haplotree.info. Retrieved 23 February 2024.
  17. ^ Chataigner, Christine (1 December 2024). "The South Caucasus from the Upper Palaeolithic to the Neolithic: Intersection of the genetic and archaeological data". Quaternary Science Reviews. 345: 109061. Bibcode:2024QSRv..34509061C. doi:10.1016/j.quascirev.2024.109061. ISSN 0277-3791. Genetic analyses, which are available only for the latter phase, reveal a new genome (Caucasus Hunter-Gatherer or CHG), that is very close to that of the ancestors of the Neolithic populations of the Zagros. For the Early Holocene (ca. 11.7–8.2 ka cal BP or 9700-6200 cal BCE), the CHG genome, which still characterises the populations of the South Caucasus, is difficult to distinguish in modelling from that of the Zagros (Iran_N).
  18. ^ Chataigner, Christine; Arimura, Makoto; Agapishvili, Tamara; Chahoud, Jwana; Koridze, Irekle; Mgeladze, Ana; Mibord, Tim; Varoutsikos, Bastien (1 September 2024). "Paravani-2, a Late Upper Palaeolithic rock-shelter site in the Javakheti highland, Southern Caucasus (Georgia)". Archaeological Research in Asia. 39: 100542. doi:10.1016/j.ara.2024.100542. ISSN 2352-2267. In fact, the CHG genome can be modelled as a mixture of three populations: 72% Western Asia_UP (ancestors of Iran_N) + 18% Caucasus_UP + 10% EHG (Eastern European Hunter-Gatherers) (Allentoft et al., 2024). [...] In the models, CHG and Iran_N often appear as interchangeable sources for Holocene populations; various names have been created to group them together, generally as Iran_N/CHG, Iran/Caucasus ancestry and Zagros/Caucasus (Wang et al., 2019; Feldman et al., 2019; Skourtanioti et al., 2020; Altınışık et al., 2022; Koptekin et al., 2023; Guarino-Vignon et al., 2023).
  19. ^ 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 (28 May 2020). "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. hdl:20.500.12154/1254. ISSN 0092-8674. PMID 32470401. Iran_C itself can be modeled as a mixture of Iran_N and Barcın_N (p = 0.365; 37% ± 3% from Barcın_N)
  20. ^ 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 (28 May 2020). "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. hdl:20.500.12154/1254. ISSN 0092-8674. PMID 32470401. 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.
  21. ^ Almarri, Mohamed A.; Haber, Marc; Lootah, Reem A.; Hallast, Pille; Al Turki, Saeed; Martin, Hilary C.; Xue, Yali; Tyler-Smith, Chris (2 September 2021). "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).
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  26. ^ Heggarty, Paul; Anderson, Cormac; Scarborough, Matthew; King, Benedict; Bouckaert, Remco; Jocz, Lechosław; Kümmel, Martin Joachim; Jügel, Thomas; Irslinger, Britta; Pooth, Roland; Liljegren, Henrik; Strand, Richard F.; Haig, Geoffrey; Macák, Martin; Kim, Ronald I. (28 July 2023). "Language trees with sampled ancestors support a hybrid model for the origin of Indo-European languages". Science. 381 (6656): eabg0818. doi:10.1126/science.abg0818. PMID 37499002.
  27. ^ Cite error: The named reference :5 was invoked but never defined (see the help page).
  28. ^ 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 (28 May 2020). "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. hdl:20.500.12154/1254. ISSN 0092-8674. PMID 32470401. 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.
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