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Early European Farmers

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Early European Farmers (EEF)[a] 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.

The earliest farmers in Anatolia derived most (80-90%) of their ancestry from the regions local hunter-gatherers, with minor Levantine and Caucasus-related ancestry.[1] The Early European Farmers moved into Europe from Anatolia through Southeast Europe from around 7,000 BC, gradually spread north and westwards, and reached Northwest Africa via the Iberian Peninsula. Genetic studies have confirmed that the later Farmers of Europe generally have also a minor contribution from Western Hunter-Gatherers (WHGs), with significant regional variation. European farmer and hunter-gatherer populations coexisted and traded in some locales, although evidence suggests that the relationship was not always peaceful. Over the course of the next 4,000 years or so, Europe was transformed into agricultural communities, with WHGs being effectively replaced across Europe. During the Chalcolithic and early Bronze Age, people who had Western Steppe Herder (WSH) ancestry moved into Europe and mingled with the EEF population; these WSH, originating from the Yamnaya culture of the Pontic steppe of Eastern Europe, probably spoke Indo-European languages. EEF ancestry is common in modern European and Northwest African populations, with EEF ancestry highest in Southern Europeans, especially Sardinians and Basque people

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

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See also: Neolithic Revolution
Spread of farming from Southwest Asia to Europe and Northwest Africa, between 9600 and 4000 BC

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][6] 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.[7]

The most common paternal haplogroup among EEFs was haplogroup G2a, while haplogroups E1b1 and R1b have also been found.[8] 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][10]

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,[11][12] suggesting the migrations involved both males and females from the steppe.[13][14]

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.[15] 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.[16]

EEF ancestry remains widespread throughout Europe, ranging from about 60% near the Mediterranean Sea (with a peak of 65% [17] 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.[18][19] EEF ancestry is also prominent in living Northwest Africans like Moroccans and Algerians.[20]

Physical appearance and allele frequency

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Reconstruction of a Neolithic farmer from Europe, Science Museum in Trento

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.[21] 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.[22]

The Early European Farmers are believed to have been mostly dark haired and dark eyed, and light skinned,[23][24] although darker than most modern Europeans.[25] A study on different EEF remains throughout Europe concluded that they had "intermediate to light skin complexion".[26] A 2024 study found that risk alleles for mood-related phenotypes are enriched in the ancestry of Neolithic farmers.[27]

Subsistence

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EEFs and their Anatolian forebears kept taurine cattle,[28] pigs,[29] sheep and goats,[30] as livestock, and planted cereal crops like wheat.[31]

Social organisation

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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.[32][33]

See also

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References

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  1. ^ a b 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. 10 (1): 1218. Bibcode:2019NatCo..10.1218F. doi:10.1038/s41467-019-09209-7. ISSN 2041-1723. PMC 6425003. PMID 30890703.
  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. doi:10.1016/j.cell.2022.04.008. ISSN 0092-8674. PMC 9166250. PMID 35561686.
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  4. ^ Mathieson et al. 2018.
  5. ^ Curry, Andrew (August 2019). "The first Europeans weren't who you might think". National Geographic. Archived from the original on 19 March 2021.
  6. ^ Spinney, Laura (1 July 2020). "When the First Farmers Arrived in Europe, Inequality Evolved". Scientific American.
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  8. ^ a b Manco, Jean (2016). Ancestral Journeys: The Peopling of Europe from the First Venturers to the Vikings (Revised and Updated ed.). Thames & Hudson. p. 98-100. ISBN 978-0-500-77290-4.
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  12. ^ 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 (15 November 2019). "Human mitochondrial DNA lineages in Iron-Age Fennoscandia suggest incipient admixture and eastern introduction of farming-related maternal ancestry". Scientific Reports. 9 (1): 16883. Bibcode:2019NatSR...916883O. doi:10.1038/s41598-019-51045-8. ISSN 2045-2322. PMC 6858343. PMID 31729399. "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)."
  13. ^ Juras et al. 2018: We identified, for the first time in ancient populations, the rare mitochondrial haplogroup X4 in two Bronze Age Catacomb culture-associated individuals. Genetic similarity analyses show close maternal genetic affinities between populations associated with both eastern and Baltic Corded Ware culture, and the Yamnaya horizon, in contrast to larger genetic differentiation between populations associated with western Corded Ware culture and the Yamnaya horizon. This indicates that females with steppe ancestry contributed to the formation of populations associated with the eastern Corded Ware culture while more local people, likely of Neolithic farmer ancestry, contributed to the formation of populations associated with western Corded Ware culture.
  14. ^ Olalde et al. 2019, pp. 1–2.
  15. ^ Goldberg et al. 2017.
  16. ^ Lazaridis, Iosif; Reich, David (5 May 2017). "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): E3873–E3874. Bibcode:2017PNAS..114E3873L. doi:10.1073/pnas.1704308114. ISSN 0027-8424. PMC 5441797. PMID 28476764.
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  19. ^ 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 (2021). The genomic history of the Aegean palatial civilizations. p. 41. OCLC 1263227362.{{cite book}}: CS1 maint: multiple names: authors list (link)
  20. ^ 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. 29 (22): 3953–3959.e4. Bibcode:2019CBio...29E3953S. doi:10.1016/j.cub.2019.09.050. PMID 31679935.
  21. ^ Martiniano et al. 2017, p. 9.
  22. ^ Mathieson et al. 2015, p. 4. "[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."
  23. ^ Reich 2018, p. 96
  24. ^ Lalueza-Fox, Carles (1 February 2022). Inequality: A Genetic History. MIT Press. p. 29. ISBN 978-0-262-04678-7. "p.29: "Physically, early farmers from Anatolia were different from those foragers; they had brown eyes but fair skin...."
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  27. ^ 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. 625 (7994): 312–320. Bibcode:2024Natur.625..312I. doi:10.1038/s41586-023-06705-1. ISSN 1476-4687. PMC 10781624. PMID 38200293.
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Bibliography

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Further reading

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Notes

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  1. ^ Sometimes called as First European Farmers, Neolithic European Farmers or Ancient Aegean Farmers
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