Early European Farmers

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 region's 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 lower levels.

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 Caucasus 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, 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 farmers of the Neolithic 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 the Bell Beaker people around 2,500 BC,^[9] who had approximately 50% WSH ancestry.^[10]

The individuals buried in Neolithic 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 the Funnelbeaker culture of southern Scandinavia 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-gatherer Pitted Ware culture; the latter carried no EEF admixture and were instead genetically similar to other European hunter-gatherers.^[12]

The most common paternal haplogroup among EEFs was haplogroup G2a, while haplogroups E1b1 and R1b have also been found.^[13] 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.^[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]

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,^[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 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.^[22]

EEF ancestry remains widespread throughout Europe, ranging from about 60% near the Mediterranean Sea (with a peak of 65% in the island of Sardinia) 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 like Moroccans and Algerians.^[26]

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.^[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, and light skinned,^[29][30] with the derived SLC24A5 being fixed in the Anatolia Neolithic,^[31] although a genetic study of Ötzi the Iceman, a Chalcolithic 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 mostly had an "intermediate to light skin complexion".^[33] A 2024 paper found that risk alleles for mood-related phenotypes are enriched in the ancestry of Neolithic farmers.^[34]

EEFs and their Anatolian forebears kept taurine cattle,^[35] pigs,^[36] sheep, and goats^[37] as livestock, and planted cereal crops like wheat.^[38]

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, who were genetically unrelated to their husbands, suggesting female exogamy.^[39][40]

A Neolithic royal buried at Newgrange 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, some settlements of the Cucuteni–Trypillia culture, such as Talianki (with a population of around 15,000) in western Ukraine, were as large as the city-states of Sumer in the Fertile 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 of state-level societies. An excavated mega-structures suggests the presence of public buildings for meetings or ceremonies.^[42]

• Neolithic Europe
• Neolithic decline
• Anatolian hunter-gatherers