Archaeogenetic name for an ancestral genetic component
Early European Farmers (EEF)[a] were a group of theAnatolian Neolithic Farmers (ANF) who brought agriculture toEurope andNorthwest Africa. The Anatolian Neolithic Farmers were an ancestral component, first identified in farmers fromAnatolia (also known as Asia Minor) in theNeolithic, and outside of Europe and Northwest Africa, they also existed inIranian Plateau,South Caucasus,Mesopotamia and theLevant. Although the spread of agriculture from theMiddle East to Europe has long been recognised through archaeology, it is only recent advances inarchaeogenetics that have confirmed that this spread was strongly correlated with amigration of these farmers, and was not just acultural exchange.
The earliest farmers in Anatolia derived most (80–90%) of their ancestry from the region'slocal hunter-gatherers, with minor Levantine and Caucasus-related ancestry.[1] The Early European Farmers moved into Europe from Anatolia throughSoutheast Europe from around 7,000 BC, gradually spread north and westwards, and reached Northwest Africa via theIberian Peninsula.Genetic studies have confirmed that the later Farmers of Europe generally have also a minor contribution fromWestern 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 theChalcolithic and earlyBronze Age, people who hadWestern Steppe Herder (WSH) ancestry moved into Europe and mingled with the EEF population; these WSH, originating from theYamnaya culture of thePontic steppe of Eastern Europe, probably spokeIndo-European languages. EEF ancestry is common in modern European and Northwest African populations, with EEF ancestry highest in Southern Europeans, especiallySardinians andBasque people.
Genetic studies demonstrate that the introduction of farming to Europe in the 7th millennium BC was associated with a mass migration of people from NorthwestAnatolia 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 theDanube (Linear Pottery culture) or the westernMediterranean (Cardial Ware). Large parts ofNorthern Europe andEastern 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, EEFs originating from the Iberian Peninsula migrated into Northwest Africa, bringing farming to the region. They were a key component in the neolithization process of the Maghreb, and intermixed with the local forager communities.[7]
The builders ofStonehenge were descendants of Neolithic farmers who migrated to the area about 6,000 years ago
The farmers of theNeolithic British Isles had entered the region through a mass migration c. 4,000 BC. They carried about 80% EEF and 20% WHG ancestry and were found to be closely related to Neolithic peoples of Iberia, which implies that they were descended from agriculturalists who had moved westwards from the Balkans along the Mediterranean coast. The arrival of farming populations led to the almost complete replacement of the native WHGs of the British Isles, who did not experience a genetic resurgence in the succeeding centuries.[8] More than 90% of Britain's Neolithic gene pool was replaced with the arrival of theBell Beaker people around 2,500 BC,[9] who had approximately 50%WSH ancestry.[10]
The individuals buried inNeolithic Ireland were found to be largely of EEF ancestry (with WHG admixture), and were closely related to peoples of Neolithic Britain and Iberia. It was found that the Neolithic peoples of Ireland had almost entirely replaced the native Irish Hunter-Gatherers through a rapid maritime colonization.[11]
The people of theFunnelbeaker culture of southernScandinavia were largely of EEF descent, with slight hunter-gatherer admixture, suggesting that the emergence of the Neolithic in Scandinavia was a result of human migration from the south. The Funnelbeakers were found to be genetically highly different from people of the neighboring hunter-gathererPitted Ware culture; the latter carried no EEF admixture and were instead genetically similar to other European hunter-gatherers.[12]
The most commonpaternal haplogroup among EEFs washaplogroup G2a, while haplogroupsE1b1 andR1b have also been found.[13] Theirmaternal haplogroups consisted mainly of West Eurasian lineages including haplogroupsH2,I, andT2, however significant numbers of central European farmers belonged to East Asian maternal lineageN9a, which is almost non-existent in modern Europeans, but common inEast Asia.[13][14][15] However, the high frequency of the East Asian mitochondrial haplogroup N9a in Neolithic cultures of the Carpathian Basin was disputed by another study.[16]
Neolithic cultures in Europe in c. 4500–4000 BC
During theChalcolithic and earlyBronze Age, the EEF-derived cultures of Europe were overwhelmed by successive migrations ofWestern Steppe Herders (WSHs) from thePontic–Caspian steppe, who carried roughly equal amounts ofEastern Hunter-Gatherer (EHG) andCaucasus Hunter-Gatherer (CHG) ancestries. These migrations led to EEFpaternal DNA lineages in Europe being almost entirely replaced with WSH-derived paternal DNA (mainly subclades of EHG-derivedR1b andR1a). EEFmaternal DNA (mainly haplogroup N) was also substantially replaced, being supplanted by steppe lineages,[17][18] suggesting the migrations involved both males and females from the steppe.[19][20]
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.[21] However, this study's resultscould not be replicated in a follow-up study by Iosif Lazaridis andDavid Reich, suggesting that the authors had mis-measured the admixture proportions of their sample.[22]
EEF ancestry remains widespread throughout Europe, ranging from about 60% near theMediterranean Sea (with a peak of 65% in the island ofSardinia) and diminishing northwards to about 10% in northern Scandinavia.[23] According to more recent studies however, 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.[24][25] EEF ancestry is also prominent in living Northwest Africans likeMoroccans andAlgerians.[26]
Reconstruction of a Neolithic farmer from Europe,Science Museum in Trento
European hunter-gatherers were muchtaller 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.[27] 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.[28]
The Early European Farmers are believed to have been mostly dark haired and dark eyed, andlight skinned,[29][30] with the derivedSLC24A5 being fixed in the Anatolia Neolithic,[31] although a genetic study ofÖtzi the Iceman, aChalcolithic mummy of EEF ancestry, found that he had a darker skin tone than contemporary southern Europeans.[32] A study on different EEF remains throughout Europe concluded that they most likely had an "intermediate to light skin complexion".[33]: s14 A 2024 paper found that risk alleles formood-related phenotypes are enriched in the ancestry of Neolithic farmers.[34]
Genetic analysis of individuals found in Neolithic tombs suggests that least some EEF peoples werepatrilineal (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, who were genetically unrelated to their husbands, suggesting femaleexogamy.[39][40]
A Neolithic royal buried atNewgrange was found to be highly inbred and possibly the product of an incestual relationship, suggesting that this community was highly socially stratified and dominated by a line of powerful "god-kings".[11]
In terms of overall size, somesettlements of the Cucuteni–Trypillia culture, such asTalianki (with a population of around 15,000) in western Ukraine, were as large as thecity-states ofSumer in theFertile Crescent, and these Eastern European settlements predate the Sumerian cities by more than half of a millennium.[41] Research indicates that the settlements had a three-level settlement hierarchy, with the possibility ofstate-level societies. An excavated mega-structures suggests the presence ofpublic buildings for meetings orceremonies.[42]
^Crabtree, Pam J.; Bogucki, Peter (25 January 2017).European Archaeology as Anthropology: Essays in Memory of Bernard Wailes. University of Pennsylvania Press. p. 55.ISBN978-1-934536-90-2.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."
^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.
^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; Mota, Barbara; Paulsson, Bettina Schulz; Halgren, Alma; Macleod, Ruairidh; Jørkov, Marie Louise Schjellerup (5 May 2022),Population Genomics of Stone Age Eurasia, pp. 2022.05.04.490594,doi:10.1101/2022.05.04.490594,hdl:2108/348203,S2CID248563160
^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.OCLC1263227362.{{cite book}}: CS1 maint: multiple names: authors list (link)
^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."
^Lalueza-Fox, Carles (1 February 2022).Inequality: A Genetic History. MIT Press. p. 29.ISBN978-0-262-04678-7. "p.29: "Physically, early farmers from Anatolia were different from those foragers; they had brown eyes but fair skin...."
^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 (supplement 6): 1842–1859.e18.doi:10.1016/j.cell.2022.04.008.ISSN0092-8674.PMC9166250.PMID35561686.We found that the vast majority of the newly sequenced EFs [European early farmers] most likely had an intermediate to light skin complexion
