Early European Farmers

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Early European Farmers (EEF) [lower-alpha 1] 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.

Contents

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.

Overview

Spread of farming from Southwest Asia to Europe and Northwest Africa, between 9600 and 4000 BC Expansion of farming in western Eurasia, 9600-4000 BCE.png
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 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 Peninsular 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 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% in the island of Sardinia) and diminishing northwards to about 10% in northern Scandinavia. [17] 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. [18] [19] EEF ancestry is also prominent in living Northwest Africans like Moroccans and Algerians. [20]

Physical appearance and allele frequency

Reconstruction of a Neolithic farmer from Europe, Science Museum in Trento Homo sapiens - Neolithic - reconstruction - MUSE.jpg
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] with the derived SLC24A5 being fixed in the Anatolia Neolithic, [25] although some farmers were slightly darker than most modern Europeans. [26] A study on different EEF remains throughout Europe concluded that they mostly had an "intermediate to light skin complexion". [27] A 2024 paper found that risk alleles for mood-related phenotypes are enriched in the ancestry of Neolithic farmers. [28]

Subsistence

EEFs and their Anatolian forebears kept taurine cattle, [29] pigs, [30] sheep, and goats [31] as livestock, and planted cereal crops like wheat. [32]

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, who were genetically unrelated to their husbands, suggesting female exogamy. [33] [34]

See also

Related Research Articles

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<span class="mw-page-title-main">Neolithic Europe</span> Era of pre-history

The European Neolithic is the period from the arrival of Neolithic technology and the associated population of Early European Farmers in Europe, c. 7000 BC until c. 2000–1700 BC. The Neolithic overlaps the Mesolithic and Bronze Age periods in Europe as cultural changes moved from the southeast to northwest at about 1 km/year – this is called the Neolithic Expansion.

<span class="mw-page-title-main">Funnelbeaker culture</span> North-central European culture around 4300–2800 BCE

The Funnel(-neck-)beaker culture, in short TRB or TBK, was an archaeological culture in north-central Europe. It developed as a technological merger of local neolithic and mesolithic techno-complexes between the lower Elbe and middle Vistula rivers. These predecessors were the (Danubian) Lengyel-influenced Stroke-ornamented ware culture (STK) groups/Late Lengyel and Baden-Boleráz in the southeast, Rössen groups in the southwest and the Ertebølle-Ellerbek groups in the north. The TRB introduced farming and husbandry as major food sources to the pottery-using hunter-gatherers north of this line.

<span class="mw-page-title-main">Yamnaya culture</span> Archaeological culture from the Pontic steppe

The Yamnaya culture or the Yamna culture, also known as the Pit Grave culture or Ochre Grave culture, is a late Copper Age to early Bronze Age archaeological culture of the region between the Southern Bug, Dniester, and Ural rivers, dating to 3300–2600 BC. It was discovered by Vasily Gorodtsov following his archaeological excavations near the Donets River in 1901–1903. Its name derives from its characteristic burial tradition: Я́мная is a Russian adjective that means 'related to pits ', as these people used to bury their dead in tumuli (kurgans) containing simple pit chambers. Research in recent years has found that Mikhaylovka, in lower Dnieper river, Ukraine, formed the Core Yamnaya culture.

<span class="mw-page-title-main">Pre-Pottery Neolithic B</span> Neolithic culture in upper Mesopotamia and the Levant c. 8800–6500 BC

Pre-Pottery Neolithic B (PPNB) is part of the Pre-Pottery Neolithic, a Neolithic culture centered in upper Mesopotamia and the Levant, dating to c. 10,800 – c. 8,500 years ago, that is, 8800–6500 BC. It was typed by British archaeologist Kathleen Kenyon during her archaeological excavations at Jericho in the West Bank, territory of Palestine.

<span class="mw-page-title-main">Cardium pottery</span> Archaeological culture

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<span class="mw-page-title-main">Globular Amphora culture</span> Archaeological culture in Central Europe

The Globular Amphora culture (GAC, German: Kugelamphoren-Kultur ; c. 3400–2800 BC, is an archaeological culture in Central Europe. Marija Gimbutas assumed an Indo-European origin, though this is contradicted by newer genetic studies that show a connection to the earlier wave of Early European Farmers rather than to Western Steppe Herders from the Ukrainian and south-western Russian steppes.

<span class="mw-page-title-main">Sredny Stog culture</span> Archaeological culture in Eastern Europe

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<span class="mw-page-title-main">Genetic history of Europe</span>

The genetic history of Europe includes information around the formation, ethnogenesis, and other DNA-specific information about populations indigenous, or living in Europe.

The genetic history of the British Isles is the subject of research within the larger field of human population genetics. It has developed in parallel with DNA testing technologies capable of identifying genetic similarities and differences between both modern and ancient populations. The conclusions of population genetics regarding the British Isles in turn draw upon and contribute to the larger field of understanding the history of the human occupation of the area, complementing work in linguistics, archaeology, history and genealogy.

<span class="mw-page-title-main">Genetic history of Italy</span>

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<span class="mw-page-title-main">Ancient North Eurasian</span> Archaeogenetic name for an ancestral genetic component

In archaeogenetics, the term Ancient North Eurasian (ANE) is the name given to an ancestral component that represents the lineage of the people of the Mal'ta–Buret' culture and populations closely related to them, such as the Upper Paleolithic individuals from Afontova Gora in Siberia. Genetic studies also revealed that the ANE are closely related to the remains of the preceding Yana Culture, which were dubbed as 'Ancient North Siberians' (ANS). Ancient North Eurasians are predominantly of West Eurasian ancestry who arrived in Siberia via the "northern route", but also derive a significant amount of their ancestry from an East Eurasian source, having arrived to Siberia via the "southern route".

<span class="mw-page-title-main">Western hunter-gatherer</span> Archaeogenetic name for an ancestral genetic component

In archaeogenetics, western hunter-gatherer is a distinct ancestral component of modern Europeans, representing descent from a population of Mesolithic hunter-gatherers who scattered over western, southern and central Europe, from the British Isles in the west to the Carpathians in the east, following the retreat of the ice sheet of the Last Glacial Maximum. It is closely associated and sometimes considered synonymous with the concept of the Villabruna cluster, named after Ripari Villabruna cave in Italy, known from the terminal Pleistocene of Europe, which is largely ancestral to later WHG populations.

<span class="mw-page-title-main">Caucasus hunter-gatherer</span> Anatomically modern human genetic lineage identified in 2015

Caucasus hunter-gatherer (CHG), also called Satsurblia cluster, is an anatomically modern human genetic lineage, first identified in a 2015 study, based on the population genetics of several modern Western Eurasian populations.

<span class="mw-page-title-main">Eastern hunter-gatherer</span> Archaeogenetic name for an ancestral genetic component

In archaeogenetics, eastern hunter-gatherer (EHG), sometimes east European hunter-gatherer or eastern European hunter-gatherer, is a distinct ancestral component that represents Mesolithic hunter-gatherers of Eastern Europe.

Basal Eurasian is a proposed lineage of anatomically modern humans with reduced, or zero, Neanderthal admixture (ancestry) compared to other ancient non-Africans. Basal Eurasians represent a sister lineage to other Eurasians and may have originated from the Southern Middle East, specifically the Arabian Peninsula, or North Africa, and are said to have contributed ancestry to various West Eurasian, South Asian, and Central Asian as well as African groups. This Basal Eurasian component is also proposed to explain the lower archaic admixture among modern West Eurasians compared to with East Eurasians, although alternatives without the need of such Basal admixture exist as well. Basal Eurasian ancestry had likely admixed into West Eurasian groups present in West Asia as early as 26,000 years ago, prior to the Last Glacial Maximum, with this ancestry being subsequently spread by later migrations, such as those of the Anatolian Neolithic Farmers into Europe during the Holocene.

Anatolian hunter-gatherer (AHG) is a distinct anatomically modern human archaeogenetic lineage, first identified in a 2019 study based on the remains of a single Epipaleolithic individual found in central Anatolia, radiocarbon dated to around 13,500 BCE. A population related to this individual was the main source of the ancestry of later Anatolian Neolithic Farmers, who along with Western Hunter Gatherers (WHG) and Ancient North Eurasians are one of the three currently known ancestral genetic contributors to present-day Europeans.

<span class="mw-page-title-main">Scandinavian hunter-gatherer</span> Archaeogenetic name for an ancestral genetic component

In archaeogenetics, the term Scandinavian hunter-gatherer (SHG) is the name given to a distinct ancestral component that represents descent from Mesolithic hunter-gatherers of Scandinavia. Genetic studies suggest that the SHGs were a mix of western hunter-gatherers (WHGs) initially populating Scandinavia from the south during the Holocene, and eastern hunter-gatherers (EHGs), who later entered Scandinavia from the north along the Norwegian coast. During the Neolithic, they admixed further with Early European Farmers (EEFs) and Western Steppe Herders (WSHs). Genetic continuity has been detected between the SHGs and members of the Pitted Ware culture (PWC), and to a certain degree, between SHGs and modern northern Europeans. The Sámi, on the other hand, have been found to be completely unrelated to the PWC.

<span class="mw-page-title-main">Western Steppe Herders</span> Archaeogenetic name for an ancestral genetic component

In archaeogenetics, the term Western Steppe Herders (WSH), or Western Steppe Pastoralists, is the name given to a distinct ancestral component first identified in individuals from the Chalcolithic steppe around the turn of the 5th millennium BC, subsequently detected in several genetically similar or directly related ancient populations including the Khvalynsk, Repin, Sredny Stog, and Yamnaya cultures, and found in substantial levels in contemporary European, Central Asian, South Asian and West Asian populations. This ancestry is often referred to as Yamnaya ancestry, Yamnaya-related ancestry, Steppe ancestry or Steppe-related ancestry.

<span class="mw-page-title-main">Genetic history of Sardinia</span>

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References

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  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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  26. ↑ Wang, Ke; Prüfer, Kay; Krause-Kyora, Ben; Childebayeva, Ainash; Schuenemann, Verena J.; Coia, Valentina; Maixner, Frank; Zink, Albert; Schiffels, Stephan; Krause, Johannes (16 August 2023). "High-coverage genome of the Tyrolean Iceman reveals unusually high Anatolian farmer ancestry". Cell Genomics. 3 (9): 100377. doi:10.1016/j.xgen.2023.100377. ISSN   2666-979X. PMC   10504632 . PMID   37719142.
  27. ↑ 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. We find that the vast majority of early farmers in our dataset had intermediate to light skin complexion
  28. ↑ 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.
  29. ↑ Scheu, Amelie; Powell, Adam; Bollongino, Ruth; Vigne, Jean-Denis; Tresset, Anne; Çakırlar, Canan; Benecke, Norbert; Burger, Joachim (December 2015). "The genetic prehistory of domesticated cattle from their origin to the spread across Europe". BMC Genetics. 16 (1): 54. doi: 10.1186/s12863-015-0203-2 . ISSN   1471-2156. PMC   4445560 . PMID   26018295.
  30. ↑ Frantz, Laurent A. F.; Haile, James; Lin, Audrey T.; Scheu, Amelie; Geörg, Christina; Benecke, Norbert; Alexander, Michelle; Linderholm, Anna; Mullin, Victoria E.; Daly, Kevin G.; Battista, Vincent M.; Price, Max; Gron, Kurt J.; Alexandri, Panoraia; Arbogast, Rose-Marie (27 August 2019). "Ancient pigs reveal a near-complete genomic turnover following their introduction to Europe". Proceedings of the National Academy of Sciences. 116 (35): 17231–17238. Bibcode:2019PNAS..11617231F. doi: 10.1073/pnas.1901169116 . ISSN   0027-8424. PMC   6717267 . PMID   31405970.
  31. ↑ Gillis, Rosalind E.; Gaastra, Jane S.; Linden, Marc Vander; Vigne, Jean-Denis (2 January 2022). "A Species Specific Investigation Into Sheep and Goat Husbandry During the Early European Neolithic". Environmental Archaeology. 27 (1): 8–19. Bibcode:2022EnvAr..27....8G. doi:10.1080/14614103.2019.1615214. ISSN   1461-4103.
  32. ↑ Brami, Maxime; Heyd, Volker (January 2011). "The origins of Europe's first farmers: The role of Hacılar and Western Anatolia, fifty years on". Praehistorische Zeitschrift. 86 (2). doi:10.1515/pz.2011.011. ISSN   0079-4848.
  33. ↑ Fowler, Chris; Olalde, Iñigo; Cummings, Vicki; Armit, Ian; Büster, Lindsey; Cuthbert, Sarah; Rohland, Nadin; Cheronet, Olivia; Pinhasi, Ron; Reich, David (27 January 2022). "A high-resolution picture of kinship practices in an Early Neolithic tomb". Nature. 601 (7894): 584–587. Bibcode:2022Natur.601..584F. doi:10.1038/s41586-021-04241-4. ISSN   0028-0836. PMC   8896835 . PMID   34937939.
  34. ↑ Seersholm, Frederik Valeur; Sjögren, Karl-Göran; Koelman, Julia; Blank, Malou; Svensson, Emma M.; Staring, Jacqueline; Fraser, Magdalena; Pinotti, Thomaz; McColl, Hugh; Gaunitz, Charleen; Ruiz-Bedoya, Tatiana; Granehäll, Lena; Villegas-Ramirez, Berenice; Fischer, Anders; Price, T. Douglas (August 2024). "Repeated plague infections across six generations of Neolithic Farmers". Nature. 632 (8023): 114–121. doi:10.1038/s41586-024-07651-2. ISSN   1476-4687. PMC   11291285 . PMID   38987589.

Bibliography

Further reading

Notes

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