Early European Farmers Explained

Early European Farmers (EEF) were a group of the Anatolian Neolithic Farmers (ANF) who brought agriculture to Europe and Northwest Africa. The Anatolian Neolithic Farmers were an ancestral component, first identified in farmers from Anatolia (also known as Asia Minor) in the Neolithic, and outside in Europe and Northwest Africa, they also existed in Iranian Plateau, South Caucasus, Mesopotamia and Levant. Although the spread of agriculture from the Middle East to Europe has long been recognised through archaeology, it is only recent advances in archaeogenetics that have confirmed that this spread was strongly correlated with a migration of these farmers, and was not just a cultural exchange.

A distinct group of the Anatolian Neolithic Farmers spread into the east of Anatolia, and left a considerable genetic legacy in Iranian Plateau, South Caucasus, Levant (during the Pre-Pottery Neolithic B) and Mesopotamia. They also have a minor role in the ethnogenesis of WSHs of Yamnaya culture.

The ANF ancestry is found in substantial levels in contemporary European, West Asian and North African populations, and also found in Central and South Asian populations (through Bactria–Margiana Archaeological Complex and Corded Ware Culture) with more lower levels.

Overview

See also: Neolithic Revolution.

Populations of the Anatolian Neolithic derived most of their ancestry from the Anatolian hunter-gatherers (AHG), with a minor geneflow from Iranian/Caucasus and Levantine related sources, suggesting that agriculture was adopted in situ by these hunter-gatherers and not spread by demic diffusion into the region.[1] Ancestors of AHGs and EEFs are believed to have split off from Western Hunter-Gatherers (WHGs) between 45kya to 26kya during the Last Glacial Maximum, and to have split from Caucasian Hunter-Gatherers (CHGs) between 25kya to 14kya.[2]

Genetic studies demonstrate that the introduction of farming to Europe in the 7th millennium BC was associated with a mass migration of people from Northwest Anatolia to Southeast Europe,[3] which resulted in the replacement of almost all (c. 98%) of the local Balkan hunter-gatherer gene pool with ancestry from Anatolian farmers.[4] [5] In the Balkans, the EEFs appear to have divided into two wings, who expanded further west into Europe along the Danube (Linear Pottery culture) or the western Mediterranean (Cardial Ware). Large parts of Northern Europe and Eastern Europe nevertheless remained unsettled by EEFs. During the Middle Neolithic there was a largely male-driven resurgence of WHG ancestry among many EEF-derived communities, leading to increasing frequencies of the hunter-gatherer paternal haplogroups among them.

Around 7,500 years ago. Iberian EEFs migrated into Northwest Africa, bringing farming to the region.[6]

The most common paternal haplogroup among EEFs was haplogroup G2a, while haplogroups E1b1 and R1b have also been found.[7] Their maternal haplogroups consisted mainly of West Eurasian lineages including haplogroups H2, I, and T2, however significant numbers of central European farmers belonged to East Asian maternal lineage N9a, which is almost non-existent in modern Europeans, but common in East Asia.[8] [9]

During the Chalcolithic and early Bronze Age, the EEF-derived cultures of Europe were overwhelmed by successive migrations of Western Steppe Herders (WSHs) from the Pontic–Caspian steppe, who carried roughly equal amounts of Eastern Hunter-Gatherer (EHG) and Caucasus Hunter-Gatherer (CHG) ancestries. These migrations led to EEF paternal DNA lineages in Europe being almost entirely replaced with WSH-derived paternal DNA (mainly subclades of EHG-derived R1b and R1a). EEF maternal DNA (mainly haplogroup N) was also substantially replaced, being supplanted by steppe lineages,[10] [11] suggesting the migrations involved both males and females from the steppe.

A 2017 study found that Bronze Age European with steppe ancestry had elevated EEF ancestry on the X chromosome, suggesting a sex bias, in which Steppe ancestry was inherited by more male than female ancestors. However, this study's results could not be replicated in a follow-up study by Iosif Lazaridis and David Reich, suggesting that the authors had mis-measured the admixture proportions of their sample.[12]

EEF ancestry remains widespread throughout Europe, ranging from about 60% near the Mediterranean Sea (with a peak of 65% [13] in the island of Sardinia) and diminishing northwards to about 10% in northern Scandinavia. According to more recent studies the highest EEF ancestry found in modern Europeans ranges from 67% to over 80% in modern Sardinians, Italians, and Iberians, with the lowest EEF ancestry found in modern Europeans ranging around 35-40% in modern Finns, Lithuanians and Latvians.[14] EEF ancestry is also prominent in living Northwest Africans like Moroccans and Algerians.[15]

Physical appearance and allele frequency

European hunter-gatherers were much taller than EEFs, and the replacement of European hunter-gatherers by EEFs resulted in a dramatic decrease in genetic height throughout Europe. During the later phases of the Neolithic, height increased among European farmers, probably due to increasing admixture with hunter-gatherers. During the Late Neolithic and Bronze Age, further reductions of EEF ancestry in Europe due to migrations of peoples with steppe-related ancestry is associated with further increases in height. High frequencies of EEF ancestry in Southern Europe might partly explain the shortness of Southern Europeans as compared to Northern Europeans, who carry increased levels of steppe-related ancestry.[16]

The Early European Farmers are believed to have been mostly dark haired and dark eyed, and light skinned,[17] although darker than most modern Europeans.[18] A study on different EEF remains throughout Europe concluded that they had "intermediate to light skin complexion".[19] A 2024 study found that risk alleles for mood-related phenotypes are enriched in the ancestry of Neolithic farmers.[20]

Subsistence

EEFs and their Anatolian forebears kept taurine cattle,[21] pigs,[22] sheep and goats,[23] as livestock, and planted cereal crops like wheat.[24]

Social organisation

Genetic analysis of individuals found in Neolithic tombs suggests that least some EEF peoples were patrilineal (tracing descent through the male line), with the tombs occupants mostly consisting of the male descendants of a single male common ancestor and their children, as well as their wives which were genetically unrelated to their husbands, suggesting female exogamy.[25] [26]

See also

Bibliography

Further reading

Notes and References

  1. Krause. Johannes. Jeong. Choongwon. Haak. Wolfgang. Posth. Cosimo. Stockhammer. Philipp W.. Mustafaoğlu. Gökhan. Fairbairn. Andrew. Bianco. Raffaela A.. Julia Gresky. 19 March 2019. Late Pleistocene human genome suggests a local origin for the first farmers of central Anatolia. Nature Communications. en. 10. 1. 1218. 10.1038/s41467-019-09209-7. 30890703. 6425003. 2019NatCo..10.1218F . 2041-1723. free.
  2. Marchi . Nina . Winkelbach . Laura . Schulz . Ilektra . Brami . Maxime . Hofmanová . Zuzana . Blöcher . Jens . Reyna-Blanco . Carlos S. . Diekmann . Yoan . Thiéry . Alexandre . Kapopoulou . Adamandia . Link . Vivian . Piuz . Valérie . Kreutzer . Susanne . Figarska . Sylwia M. . Ganiatsou . Elissavet . May 2022 . The genomic origins of the world's first farmers . Cell . 185 . 11 . 1842–1859.e18 . 10.1016/j.cell.2022.04.008 . 0092-8674 . 9166250 . 35561686.
  3. 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 . 8 August 2016 . Genomic insights into the origin of farming in the ancient Near East . Nature . en . 536 . 7617 . 419–424 . 10.1038/nature19310 . 27459054. 5003663 . 2016Natur.536..419L .
  4. News: Andrew . Curry . The first Europeans weren't who you might think . https://web.archive.org/web/20210319032852/https://www.nationalgeographic.com/culture/article/first-europeans-immigrants-genetic-testing-feature . dead . 19 March 2021 . National Geographic . August 2019.
  5. News: Laura . Spinney . When the First Farmers Arrived in Europe, Inequality Evolved . Scientific American . 1 July 2020.
  6. Simões . Luciana G. . Günther . Torsten . Martínez-Sánchez . Rafael M. . Vera-Rodríguez . Juan Carlos . Iriarte . Eneko . Rodríguez-Varela . Ricardo . Bokbot . Youssef . Valdiosera . Cristina . Jakobsson . Mattias . 2023-06-15 . Northwest African Neolithic initiated by migrants from Iberia and Levant . Nature . en . 618 . 7965 . 550–556 . 10.1038/s41586-023-06166-6 . 0028-0836 . 10266975 . 37286608. 2023Natur.618..550S .
  7. Book: Manco . Jean . Ancestral Journeys: The Peopling of Europe from the First Venturers to the Vikings . 2016 . Thames & Hudson . 978-0-500-77290-4 . 98-100 . Revised and Updated .
  8. Guba . Zsuzsanna . Hadadi . Éva . Major . Ágnes . Furka . Tünde . Juhász . Emese . Koós . Judit . Nagy . Károly . Zeke . Tamás . HVS-I polymorphism screening of ancient human mitochondrial DNA provides evidence for N9a discontinuity and East Asian haplogroups in the Neolithic Hungary . Journal of Human Genetics . November 2011 . 56 . 11 . 784–796 . 10.1038/jhg.2011.103 . 21918529 . 20827921 . en . 1435-232X. free .
  9. Derenko . Miroslava . Malyarchuk . Boris . Grzybowski . Tomasz . Denisova . Galina . Rogalla . Urszula . Perkova . Maria . Dambueva . Irina . Zakharov . Ilia . Origin and Post-Glacial Dispersal of Mitochondrial DNA Haplogroups C and D in Northern Asia . PLOS ONE . 21 December 2010 . 5 . 12 . e15214 . 10.1371/journal.pone.0015214 . 21203537 . 3006427 . en . 1932-6203 . free . 2010PLoSO...515214D .
  10. Book: Crabtree . Pam J. . Bogucki . Peter . European Archaeology as Anthropology: Essays in Memory of Bernard Wailes . 25 January 2017 . University of Pennsylvania Press . 978-1-934536-90-2 . 55 . en. p.55: "In addition, uniparental markers changed suddenly as mtDNA N1a and Y haplogroup G2a, which had been very common in the EEF agricultural population, were replaced by Y haplogroups R1a and R1b and by a variety of mtDNA haplogroups typical of the Steppe Yamnaya population. The uniparental markers show that the migrants included both men and women from the steppes."
  11. Översti . Sanni . Majander . Kerttu . Salmela . Elina . Salo . Kati . Arppe . Laura . Belskiy . Stanislav . Etu-Sihvola . Heli . Laakso . Ville . Mikkola . Esa . Pfrengle . Saskia . Putkonen . Mikko . Taavitsainen . Jussi-Pekka . Vuoristo . Katja . Wessman . Anna . Sajantila . Antti . Oinonen . Markku . Haak . Wolfgang . Schuenemann . Verena J. . Krause . Johannes . Palo . Jukka U. . Onkamo . Päivi . Human mitochondrial DNA lineages in Iron-Age Fennoscandia suggest incipient admixture and eastern introduction of farming-related maternal ancestry . Scientific Reports . 15 November 2019 . 9 . 1 . 16883 . 10.1038/s41598-019-51045-8 . 31729399 . 6858343 . 2019NatSR...916883O . en . 2045-2322. "The subsequent spread of Yamnaya-related people and Corded Ware Culture in the late Neolithic and Bronze Age were accompanied with the increase of haplogroups I, U2 and T1 in Europe (See8 and references therein)."
  12. Lazaridis . Iosif . Reich . David . Failure to replicate a genetic signal for sex bias in the steppe migration into central Europe . Proceedings of the National Academy of Sciences . 114 . 20 . 5 May 2017 . E3873–E3874 . 0027-8424 . 10.1073/pnas.1704308114 . 28476764 . 5441797 . free . 2017PNAS..114E3873L .
  13. 10.1038/s41559-020-1102-0 . The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean . 2020 . Fernandes . Daniel M. . Mittnik . Alissa . Olalde . Iñigo . Lazaridis . Iosif . Cheronet . Olivia . Rohland . Nadin . Mallick . Swapan . Bernardos . Rebecca . Broomandkhoshbacht . Nasreen . Carlsson . Jens . Culleton . Brendan J. . Ferry . Matthew . Gamarra . Beatriz . Lari . Martina . Mah . Matthew . Michel . Megan . Modi . Alessandra . Novak . Mario . Oppenheimer . Jonas . Sirak . Kendra A. . Stewardson . Kristin . Mandl . Kirsten . Schattke . Constanze . Özdoğan . Kadir T. . Lucci . Michaela . Gasperetti . Gabriella . Candilio . Francesca . Salis . Gianfranca . Vai . Stefania . Camarós . Edgard . Nature Ecology & Evolution . 4 . 3 . 334–345 . 32094539 . 7080320 . 2020NatEE...4..334F . 1 .
  14. Book: Christina, Clemente, Florian Unterlaender, Martina Dolgova, Olga Amorim, Carlos Eduardo G. Coroado-Santos, Francisco Neuenschwander, Samuel Ganiatsou, Elissavet Davalos, Diana I. Cruz Anchieri, Lucas Michaud, Frederic Winkelbach, Laura Bloecher, Jens Cardenas, Yami Ommar Arizmendi da Mota, Barbara Sousa Kalliga, Eleni Souleles, Angelos Kontopoulos, Ioannis Karamitrou-Mentessidi, Georgia Philaniotou, Olga Sampson, Adamantios Theodorou, Dimitra Tsipopoulou, Metaxia Akamatis, Ioannis Halstead, Paul Kotsakis, Kostas Urem-Kotsou, Dushka Panagiotopoulos, Diamantis Ziota, Christina Triantaphyllou, Sevasti Delaneau, Olivier Jensen, Jeffrey D. Victor Moreno-Mayar, J. Burger, Joachim Sousa, Vitor C. Lao, Oscar Malaspinas, Anna-Sapfo Papageorgopoulou . The genomic history of the Aegean palatial civilizations . 2021 . 41 . 1263227362.
  15. Serra-Vidal . Gerard . Lucas-Sanchez . Marcel . Fadhlaoui-Zid . Karima . Bekada . Asmahan . Zalloua . Pierre . Comas . David . November 2019 . Heterogeneity in Palaeolithic Population Continuity and Neolithic Expansion in North Africa . Current Biology . en . 29 . 22 . 3953–3959.e4 . 10.1016/j.cub.2019.09.050. 31679935 . 2019CBio...29E3953S .
  16. . "[R]esults suggest that the modern South-North gradient in height across Europe is due to both increased steppe ancestry in northern populations, and selection for decreased height in Early Neolithic migrants to southern Europe."
  17. Book: Lalueza-Fox . Carles . Inequality: A Genetic History . 1 February 2022 . MIT Press . 978-0-262-04678-7 . 29 . en. "p.29: "Physically, early farmers from Anatolia were different from those foragers; they had brown eyes but fair skin...."
  18. Wang . Ke . Prüfer . Kay . Krause-Kyora . Ben . Childebayeva . Ainash . Schuenemann . Verena J. . Coia . Valentina . Maixner . Frank . Zink . Albert . Schiffels . Stephan . Krause . Johannes . 2023-08-16 . High-coverage genome of the Tyrolean Iceman reveals unusually high Anatolian farmer ancestry . Cell Genomics . 3 . 9 . 100377 . 10.1016/j.xgen.2023.100377 . 37719142 . 10504632 . 2666-979X.
  19. Marchi . Nina . Winkelbach . Laura . Schulz . Ilektra . Brami . Maxime . Hofmanová . Zuzana . Blöcher . Jens . Reyna-Blanco . Carlos S. . Diekmann . Yoan . Thiéry . Alexandre . Kapopoulou . Adamandia . Link . Vivian . Piuz . Valérie . Kreutzer . Susanne . Figarska . Sylwia M. . Ganiatsou . Elissavet . May 2022 . The genomic origins of the world's first farmers . Cell . 185 . 11 . 1842–1859.e18 . 10.1016/j.cell.2022.04.008 . 0092-8674 . 9166250 . 35561686 . We find that the vast majority of early farmers in our dataset had intermediate to light skin complexion.
  20. Irving-Pease . Evan K. . Refoyo-Martínez . Alba . Barrie . William . Ingason . Andrés . Pearson . Alice . Fischer . Anders . Sjögren . Karl-Göran . Halgren . Alma S. . Macleod . Ruairidh . Demeter . Fabrice . Henriksen . Rasmus A. . Vimala . Tharsika . McColl . Hugh . Vaughn . Andrew H. . Speidel . Leo . January 2024 . The selection landscape and genetic legacy of ancient Eurasians . Nature . en . 625 . 7994 . 312–320 . 10.1038/s41586-023-06705-1 . 38200293 . 10781624 . 2024Natur.625..312I . 1476-4687.
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