Caucasus hunter-gatherer: Difference between revisions

{{Use dmy dates|date=AprilAugust 2024}}

'''Caucasus hunter-gatherer''' ('''CHG'''), also called '''Satsurblia cluster''',{{sfn|Fu|Posth|Hajdinjak|Petr|2016}}<ref>{{cite journal |author1=Eisenmann, S. |author2= Bánffy, E. |author3=van Dommelen, P. |display-authors=etal |year=2018 |title=Reconciling material cultures in archaeology with genetic data: The nomenclature of clusters emerging from archaeogenomic analysis |journal=Scientific Reports |volume=8 |issue= 1 |pages= 13003 |doi=10.1038/s41598-018-31123-z |pmid= 30158639 |pmc= 6115390 |bibcode= 2018NatSR...813003E }}</ref> is an [[anatomically modern human]] [[human genetics|genetic lineage]], first identified in a 2015 study,{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}{{sfn|Fu|Posth|Hajdinjak|Petr|2016}} based on the [[population genetics]] of several modern [[Eurasia|Western Eurasian]] ([[Europe]]an, [[Caucasus|Caucasian]] and [[Near Eastern]]) populations.<ref name="bbcnov16"/><ref name="Dutchen 2016">{{cite web|last1=Dutchen|first1=Stephanie|title=History on Ice|url=http://hms.harvard.edu/news/history-ice|publisher=[[Harvard Medical School]]|access-date=11 May 2016|date=2 May 2016}}</ref>

It represents an ancestry maximised in some [[Upper Paleolithic]] and [[Mesolithic]] hunter-gatherer groups in the [[Caucasus]]. These groups are also very closely related to Mesolithic hunter-gatherers and [[Neolithic]] farmers in the [[Iranian Plateau]], who are sometimes included within the CHG group.<ref>{{Cite journal |lastlast1=Lazaridis |firstfirst1=Iosif |last2=Alpaslan-Roodenberg |first2=Songül |last3=Acar |first3=Ayşe |last4=Açıkkol |first4=Ayşen |last5=Agelarakis |first5=Anagnostis |last6=Aghikyan |first6=Levon |last7=Akyüz |first7=Uğur |last8=Andreeva |first8=Desislava |last9=Andrijašević |first9=Gojko |last10=Antonović |first10=Dragana |last11=Armit |first11=Ian |last12=Atmaca |first12=Alper |last13=Avetisyan |first13=Pavel |last14=Aytek |first14=Ahmet İhsan |last15=Bacvarov |first15=Krum |date=26 August 2022-08-26 |title=The genetic history of the Southern Arc: A bridge between West Asia and Europe |url=https://www.science.org/doi/10.1126/science.abm4247 |journal=Science |language=en |volume=377 |issue=6609 |pages=eabm4247 |doi=10.1126/science.abm4247 |issn=0036-8075 |pmc=PMC1006455310064553 |pmid=36007055}}</ref><ref>{{Cite journal |lastlast1=Wang |firstfirst1=Chuan-Chao |last2=Reinhold |first2=Sabine |last3=Kalmykov |first3=Alexey |last4=Wissgott |first4=Antje |last5=Brandt |first5=Guido |last6=Jeong |first6=Choongwon |last7=Cheronet |first7=Olivia |last8=Ferry |first8=Matthew |last9=Harney |first9=Eadaoin |last10=Keating |first10=Denise |last11=Mallick |first11=Swapan |last12=Rohland |first12=Nadin |last13=Stewardson |first13=Kristin |last14=Kantorovich |first14=Anatoly R. |last15=Maslov |first15=Vladimir E. |date=4 February 2019-02-04 |title=Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions |url=https://www.nature.com/articles/s41467-018-08220-8 |journal=Nature Communications |language=en |volume=10 |issue=1 |page=590 |doi=10.1038/s41467-018-08220-8 |issn=2041-1723 |pmc=PMC63601916360191 |pmid=30713341|bibcode=2019NatCo..10..590W }}</ref> Ancestry that is closely related to CHG-Iranian Neolithic farmers is also known from further east, including from the [[Bactria–Margiana Archaeological Complex]] and the [[Indus Valley Civilisation|Harappan/Indus Valley Civilisation]].<ref name=":2">{{Cite journal |lastlast1=Shinde |firstfirst1=Vasant |last2=Narasimhan |first2=Vagheesh M. |last3=Rohland |first3=Nadin |last4=Mallick |first4=Swapan |last5=Mah |first5=Matthew |last6=Lipson |first6=Mark |last7=Nakatsuka |first7=Nathan |last8=Adamski |first8=Nicole |last9=Broomandkhoshbacht |first9=Nasreen |last10=Ferry |first10=Matthew |last11=Lawson |first11=Ann Marie |last12=Michel |first12=Megan |last13=Oppenheimer |first13=Jonas |last14=Stewardson |first14=Kristin |last15=Jadhav |first15=Nilesh |date=October 2019 |title=An Ancient Harappan Genome Lacks Ancestry from Steppe Pastoralists or Iranian Farmers |url=https://linkinghub.elsevier.com/retrieve/pii/S0092867419309675 |journal=Cell |language=en |volume=179 |issue=3 |pages=729–735.e10 |doi=10.1016/j.cell.2019.08.048 |pmc=6800651 |pmid=31495572}}</ref><ref Itname=":3">{{Cite alongjournal with|last1=Narasimhan the|first1=Vagheesh M. |last2=Patterson |first2=Nick |last3=Moorjani |first3=Priya |last4=Rohland |first4=Nadin |last5=Bernardos |first5=Rebecca |last6=Mallick |first6=Swapan |last7=Lazaridis |first7=Iosif |last8=Nakatsuka |first8=Nathan |last9=Olalde |first9=Iñigo |last10=Lipson |first10=Mark |last11=Kim |first11=Alexander M. |last12=Olivieri |first12=Luca M. |last13=Coppa |first13=Alfredo |last14=Vidale |first14=Massimo |last15=Mallory |first15=James |date=6 September 2019 |title=The formation of human populations in South and Central Asia |journal=Science |language=en |volume=365 |issue=6457 |doi=10.1126/science.aat7487 |issn=0036-8075 |pmc=6822619 |pmid=31488661}}</ref> Caucasus hunter-gatherers and [[Eastern hunter-gatherer|Eastern Hunter Gather]] ancestrys are ancestral in roughly equal proportions to the [[Western Steppe Herder]]s (WSH) ancestry, whichwho waswere widely spread across Europe and Asia beginning during the [[Chalcolithic]].

| caption1 = Genetic structure of ancient Europe. Caucasus hunter-gatherers are represented by the Satsurbila and Kotias specimens.{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}

| caption2 = Genetic affinity of modern populations to the ancient Kotias specimen.{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}

| caption3 = Admixture graph of deep Eurasian lineages (Allentoft et al. 2024).<ref name="Allentoft_2024"/>

The CHG lineage is suggested to have diverged from the ancestor of [[Western Hunter-Gatherers]] (WHGs) probably during the [[Last Glacial Maximum]] (sometimessometime between 45,000 to 26,000 years ago).<ref>{{cite journal |title='Fourth strand' of European ancestry originated with hunter-gatherers isolated by Ice Age |journal=University of Cambridge |date=16 November 2015 |url=https://www.cam.ac.uk/research/news/fourth-strand-of-european-ancestry-originated-with-hunter-gatherers-isolated-by-ice-age |language=en|quote=By reading the DNA, the researchers were able to show that the lineage of this fourth Caucasus hunter-gatherer strand diverged from the western hunter-gatherers just after the expansion of anatomically modern humans into Europe from Africa.}}</ref> They further separated from the [[Anatolian hunter-gatherers|Anatolian hunter-gatherer]] (AHG) lineage later, suggested to around 25,000 years ago during the late LGM period.{{sfn|Fu|Posth|Hajdinjak|Petr|2016}}<ref>{{Cite journal |last1=Marchi |first1=Nina |last2=Winkelbach |first2=Laura |last3=Schulz |first3=Ilektra |last4=Brami |first4=Maxime |last5=Hofmanová |first5=Zuzana |last6=Blöcher |first6=Jens |last7=Reyna-Blanco |first7=Carlos S. |last8=Diekmann |first8=Yoan |last9=Thiéry |first9=Alexandre |last10=Kapopoulou |first10=Adamandia |last11=Link |first11=Vivian |last12=Piuz |first12=Valérie |last13=Kreutzer |first13=Susanne |last14=Figarska |first14=Sylwia M. |last15=Ganiatsou |first15=Elissavet |date=May 2022 |title=The genomic origins of the world's first farmers |url=|journal=Cell |volume=185 |issue=11 |pages=1842–1859.e18 |doi=10.1016/j.cell.2022.04.008 |issn=0092-8674 |pmc=9166250 |pmid=35561686}}</ref> The Caucasus hunter-gatherers managed to survive in isolation since the late LGM period as a distinct population, and display high genetic affinities to Mesolithic and Neolithic populations on the Iranian plateau, such as Neolithic specimens found in [[Ganj Dareh]]. The CHG display higher genetic affinities to European and Anatolian groups than Iranian hunter-gatherers do, suggesting a possible cline and geneflow into the CHG and less into Mesolithic and Neolithic Iranian groups.{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}<ref>{{Cite journal |last1=Gallego-Llorente |first1=M. |last2=Connell |first2=S. |last3=Jones |first3=E. R. |last4=Merrett |first4=D. C. |last5=Jeon |first5=Y. |last6=Eriksson |first6=A. |last7=Siska |first7=V. |last8=Gamba |first8=C. |last9=Meiklejohn |first9=C. |last10=Beyer |first10=R. |last11=Jeon |first11=S. |last12=Cho |first12=Y. S. |last13=Hofreiter |first13=M. |last14=Bhak |first14=J. |last15=Manica |first15=A. |date=9 August 2016 |title=The genetics of an early Neolithic pastoralist from the Zagros, Iran |journal=Scientific Reports |language=en |volume=6 |issue=1 |pages=31326 |doi=10.1038/srep31326 |pmid=27502179 |pmc=4977546 |bibcode=2016NatSR...631326G |issn=2045-2322}}</ref>

TheAccording to one model, the Mesolithic/Neolithic Iranian lineage basal to the Caucasus hunter-gatherers are inferred to derive significant amounts of their ancestry from [[Basal Eurasian]] ({{Circa|38–48%}}), with the remainder ancestry being closer to [[Ancient North Eurasian]]s or [[Eastern European Hunter-Gatherer]] (ANE/EHG; {{Circa|52–62%}}). The CHG displayed an additional ANE-like component ({{Circa|10%}}) than the Neolithic Iranians do, suggesting they may have stood in continuous contact with [[Eastern Hunter-Gatherer]]s to their North. The CHG also carry around 20% additional Paleolithic Caucasus/Anatolian ancestry. Lazaridis et. al (2016) models the CHG as a mixture of Neolithic Iranians, [[Western Hunter-Gatherer]]s and Eastern Hunter-Gatherers. In addition, CHG cluster with early Iranian farmers, who significantly do not share alleles with early Levantine farmers.{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}{{sfn|Lazaridis|Nadel|Rollefson|Merrett|2016}}<ref>{{Cite journal |last1=Almarri |first1=Mohamed A. |last2=Haber |first2=Marc |last3=Lootah |first3=Reem A. |last4=Hallast |first4=Pille |last5=Al Turki |first5=Saeed |last6=Martin |first6=Hilary C. |last7=Xue |first7=Yali |last8=Tyler-Smith |first8=Chris |date=September 2021 |title=The genomic history of the Middle East |url=https://doi.org/10.1016/j.cell.2021.07.013 |journal=Cell |volume=184 |issue=18 |pages=4612–4625.e14 |doi=10.1016/j.cell.2021.07.013 |issn=0092-8674 |pmc=8445022 |pmid=34352227}}</ref>

At the beginning of the [[Neolithic]], at {{Circa|8000 BC}}, they were probably distributed across western Iran and the Caucasus,{{sfn|Anthony|2009b|p=29}} and people similar to northern Caucasus and Iranian plateau hunter-gatherers arrived before 6000 BC in Pakistan and north-west India.{{sfn|Narasimhan|Patterson|Moorjani|Rohland|2019|p<ref name=11}}":3" /> A roughly equal merger between the CHG and [[Eastern Hunter-Gatherer]]s in the [[Pontic–Caspian steppe]] resulted in the formation of the [[Western Steppe Herders]] (WSHs). The WSHs formed the [[Yamnaya culture]] and subsequently expanded massively throughout Europe during the [[Late Neolithic]] and [[Early Bronze Age]] {{Circa|3000—2000 BC}}.{{sfn|Jeong|Balanovsky|Lukianova|Kahbatkyzy|2019}}

Caucasus hunter gatherer/Iranian-like ancestry, was first reported as maximized in hunter-gatherers from the [[South Caucasus]] and early herders/farmers in [[Azerbaijan (Iran)|northwestern Iran]], particularly the [[Zagros Mountains|Zagros]], hence the label "'''CHG/Iranian'''”.<ref>{{Cite journal |last1=Heggarty |first1=Paul |last2=Anderson |first2=Cormac |last3=Scarborough |first3=Matthew |last4=King |first4=Benedict |last5=Bouckaert |first5=Remco |last6=Jocz |first6=Lechosław |last7=Kümmel |first7=Martin Joachim |last8=Jügel |first8=Thomas |last9=Irslinger |first9=Britta |last10=Pooth |first10=Roland |last11=Liljegren |first11=Henrik |last12=Strand |first12=Richard F. |last13=Haig |first13=Geoffrey |last14=Macák |first14=Martin |last15=Kim |first15=Ronald I. |date=2023-07-28 July 2023 |title=Language trees with sampled ancestors support a hybrid model for the origin of Indo-European languages |url=https://www.science.org/doi/10.1126/science.abg0818 |journal=Science |language=en |volume=381 |issue=6656 |pages=eabg0818 |doi=10.1126/science.abg0818 |pmid=37499002 |issn=0036-8075|hdl=10234/204329 |hdl-access=free }}</ref>

According to Narasimhan et al. (2019) Iranian farmer related people arrived before 6000 BCE in Pakistan and north-west India, before the advent of farming in northern India. They suggest the possibility that this "Iranian farmer–related ancestry [...] was [also] characteristic of northern Caucasus and Iranian plateau hunter-gatherers."{{sfn|Narasimhan|Patterson|Moorjani|Rohland|2019|p<ref name=11}}":3" />

[[File:Admixture_proportions_of_Yamnaya_populations.png|thumb|Admixture proportions of Yamnaya populations: they combined as [[Eastern Hunter Gatherer]] (<small>{{Colorsample|#00BFFF|0.6}}</small> EHG) and [[Caucasian Hunter-Gatherer]] (<small>{{Colorsample|#228B22|0.6}}</small> CHG), and the small proportions of Anatolian Farmer (<small>{{Colorsample|#FFA500|0.6}}</small> Anatolian Neolithic) and Western Hunter Gatherer (<small>{{Colorsample|#0000FF|0.6}}</small> WHG) ancestry.<ref name=":1">{{cite journal |last1=Wang |first1=Chuan-Chao |last2=Reinhold |first2=Sabine |last3=Kalmykov |first3=Alexey |date=February 4, February 2019 |title=Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions |journal=Nature Communications |language=en |volume=10 |issue=1 |pages=590 |bibcode=2019NatCo..10..590W |doi=10.1038/s41467-018-08220-8 |issn=2041-1723 |pmc=6360191 |pmid=30713341}}</ref>]]

According to Jones ''et al''. (2015), Caucasus hunter-gatherer (CHG) "genomes significantly contributed to the Yamnaya steppe herders who migrated into Europe ~3,000 BCE, supporting a formative Caucasus influence on this important Early [[Bronze Age]] culture. CHG left their imprint on modern populations from the [[Caucasus]] and also [[Central Asia|Central]] and [[South Asia]] possibly correlating with the arrival of [[Indo-Aryan language]]s."<ref>{{harvnb|Jones|Gonzalez-Fortes|Connell|Siska|2015}}: "Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers ~45 kya, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers ~25 kya, around the Last Glacial Maximum."</ref> For example, about 50%-7050–70% of Armenian ancestry is derived from CHG, persisting from Neolithic times to the present.<ref>{{Cite journal |last1=Dergachev |first1=Valentin |last2=Shephard |first2=Henry |last3=Sirbu |first3=Ghenadie |last4=Szécsényi-Nagy |first4=Anna |date=2022 |title=The genetic history of the Southern Arc: A bridge between West Asia and Europe |url=https://www.researchgate.net/publication/363157308 |journal=Science |volume=377 |issue=6609 |pages=982–987 |via=ResearchGate}}</ref> Wang et al. (2018) analysed genetic data of the North Caucasus of fossils dated between the 4th and 1st millennia BC and found correlation with modern groups of the South Caucasus, concluding that "unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement".{{sfn|Wang|Reinhold|Kalmykov|Wissgott|2018}} According to Allentoft ''et al.'' (2024), The arrival and admixture of CHG with Caspian steppe cultures is dated to about 7,300-years-old, which is seen in two ancient samples from Golubaya Krinitsa with 18-2418–24% admixture.<ref name="Allentoft_2024">{{Cite journal |last1=Allentoft |first1=Morten E. |last2=Sikora |first2=Martin |last3=Refoyo-Martínez |first3=Alba |last4=Irving-Pease |first4=Evan K. |last5=Fischer |first5=Anders |last6=Barrie |first6=William |last7=Ingason |first7=Andrés |last8=Stenderup |first8=Jesper |last9=Sjögren |first9=Karl-Göran |last10=Pearson |first10=Alice |last11=Sousa da Mota |first11=Bárbara |last12=Schulz Paulsson |first12=Bettina |last13=Halgren |first13=Alma |last14=Macleod |first14=Ruairidh |last15=Jørkov |first15=Marie Louise Schjellerup |date=January 2024 |title=Population genomics of post-glacial western Eurasia |url=https://www.nature.com/articles/s41586-023-06865-0 |journal=Nature |language=en |volume=625 |issue=7994 |pages=301–311 |doi=10.1038/s41586-023-06865-0 |pmid=38200295 |pmc=10781627 |bibcode=2024Natur.625..301A |issn=1476-4687|hdl=10261/360968 |hdl-access=free }}</ref>

Genetic analysis shows that Iranian-related ancestry, which was widespread in the Aegean by the Middle Bronze Age in association with the Minoan and Mycenaean cultures, had also spread as far west as Sicily in substantial proportion at least by the time of the Mycenaeans. One possibility is this ancestry spread west along with the Mycenaean cultural expansion.<ref>{{Cite journal |last1=Fernandes |first1=Daniel M. |last2=Mittnik |first2=Alissa |last3=Olalde |first3=Iñigo |last4=Lazaridis |first4=Iosif |last5=Cheronet |first5=Olivia |last6=Rohland |first6=Nadin |last7=Mallick |first7=Swapan |last8=Bernardos |first8=Rebecca |last9=Broomandkhoshbacht |first9=Nasreen |last10=Carlsson |first10=Jens |last11=Culleton |first11=Brendan J. |last12=Ferry |first12=Matthew |last13=Gamarra |first13=Beatriz |last14=Lari |first14=Martina |last15=Mah |first15=Matthew |date=March 2020 |title=The Spread of Steppe and Iranian Related Ancestry in the Islands of the Western Mediterranean |journal=Nature Ecology & Evolution |volume=4 |issue=3 |pages=334–345 |doi=10.1038/s41559-020-1102-0 |issn=2397-334X |pmc=7080320 |pmid=32094539|bibcode=2020NatEE...4..334F }}</ref> An arrival of the CHG-related component in Southern Italy from the Southern part of the Balkan Peninsula, including the Peloponnese, is compatible with the identification of genetic corridors linking the two regions and the presence of Southern European ancient signatures in Italy.<ref>{{Cite journal |last1=Raveane |first1=A. |last2=Aneli |first2=S. |last3=Montinaro |first3=F. |last4=Athanasiadis |first4=G. |last5=Barlera |first5=S. |last6=Birolo |first6=G. |last7=Boncoraglio |first7=G. |last8=Di Blasio |first8=A. M. |last9=Di Gaetano |first9=C. |last10=Pagani |first10=L. |last11=Parolo |first11=S. |last12=Paschou |first12=P. |last13=Piazza |first13=A. |last14=Stamatoyannopoulos |first14=G. |last15=Angius |first15=A. |date=4 September 2019-09-04 |title=Population structure of modern-day Italians reveals patterns of ancient and archaic ancestries in Southern Europe |journal=Science Advances |volume=5 |issue=9 |pages=eaaw3492 |doi=10.1126/sciadv.aaw3492 |issn=2375-2548 |pmc=6726452 |pmid=31517044|bibcode=2019SciA....5.3492R }}</ref> Collected data from Iron Age individuals dating from 900 to 200 BCE (including the Republican period) group shows a clear ancestry shift from the Copper Age, interpreted by ADMIXTURE as the addition of a Steppe-related ancestry component, and an increase in the Neolithic-Iranian component.<ref>{{Cite journal |last1=Antonio |first1=Margaret L. |last2=Gao |first2=Ziyue |last3=Moots |first3=Hannah M. |last4=Lucci |first4=Michaela |last5=Candilio |first5=Francesca |last6=Sawyer |first6=Susanna |last7=Oberreiter |first7=Victoria |last8=Calderon |first8=Diego |last9=Devitofranceschi |first9=Katharina |last10=Aikens |first10=Rachael C. |last11=Aneli |first11=Serena |last12=Bartoli |first12=Fulvio |last13=Bedini |first13=Alessandro |last14=Cheronet |first14=Olivia |last15=Cotter |first15=Daniel J. |date=8 November 2019-11-08 |title=Ancient Rome: A genetic crossroads of Europe and the Mediterranean |journal=Science |volume=366 |issue=6466 |pages=708–714 |doi=10.1126/science.aay6826 |issn=0036-8075 |pmc=7093155 |pmid=31699931|bibcode=2019Sci...366..708A }}</ref>