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Early European Farmers (EEF), First European Farmers (FEF), Neolithic European Farmers, Ancient Aegean Farmers, or Anatolian Neolithic Farmers (ANF) are names used to describe a distinct group of early
Neolithic The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several part ...
farmers who brought agriculture to Europe. Although the spread of agriculture from the Middle East to Europe has long been recognised through archaeology, it is only recent advances in
archaeogenetics Archaeogenetics is the study of ancient DNA using various molecular genetic methods and DNA resources. This form of genetic analysis can be applied to human, animal, and plant specimens. Ancient DNA can be extracted from various fossilized specime ...
that have confirmed that this spread was strongly correlated with a migration of these farmers, and was not just a cultural exchange. The Early European Farmers moved into Europe through the Balkans starting some 9,000 years ago, and gradually spread north and westwards. Genetic studies have confirmed that they interbred somewhat with the existing inhabitants of the region, the
Western Hunter-Gatherer In archaeogenetics, the term Western Hunter-Gatherer (WHG), West European Hunter-Gatherer or Western European Hunter-Gatherer names a distinct ancestral component of modern Europeans, representing descent from a population of Mesolithic hunter-g ...
s (WHGs) and they 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, and WHGs were displaced to the margins. During the Chalcolithic and early
Bronze Age The Bronze Age is a historic period, lasting approximately from 3300 BC to 1200 BC, characterized by the use of bronze, the presence of writing in some areas, and other early features of urban civilization. The Bronze Age is the second prin ...
, the Early European Farmer cultures were overwhelmed by new migrations from a group identified as
Western Steppe Herders 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 Eneolithic steppe around the turn of the 5th millennium BCE, ...
(WSHs) or Yamnaya. Yamnaya culture is associated with the spread of
Indo-European languages The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, Du ...
, and so the group are sometimes known as
Proto-Indo-Europeans The Proto-Indo-Europeans are a hypothetical prehistoric population of Eurasia who spoke Proto-Indo-European (PIE), the ancestor of the Indo-European languages according to linguistic reconstruction. Knowledge of them comes chiefly from ...
. Once again the populations interbred and EEF ancestry is common in modern European populations.


Overview

It has been discovered that populations of the Anatolian Neolithic derived a significant portion of their ancestry from the
Anatolian hunter-gatherers Epipaleolithic that inhabited central around 13,642-13,073 BCE. This population constitutes one of the three currently known ancestral genetic contributions of the present-day Europeans: Western Hunter Gatherers (WHG), Anatolian Neolithic Farmers ...
(AHG), suggesting that
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
was adopted in site by these hunter-gatherers and not spread by
demic diffusion Demic diffusion, as opposed to trans-cultural diffusion, is a demographic term referring to a migratory model, developed by Luigi Luca Cavalli-Sforza, of population diffusion into and across an area that had been previously uninhabited by that gro ...
into the region. Ancestors of AHGs and EEFs are believed to have split off from
Western Hunter-Gatherer In archaeogenetics, the term Western Hunter-Gatherer (WHG), West European Hunter-Gatherer or Western European Hunter-Gatherer names a distinct ancestral component of modern Europeans, representing descent from a population of Mesolithic hunter-g ...
s (WHGs) around 43,000 BC, and to have split from Caucasian Hunter-Gatherers (CHGs) around 23,000 BC. They appear to have migrated from Anatolia to the Balkans in large numbers during the 7th millennium BC, where they almost completely replaced the WHGs. The
Y-DNA The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abs ...
of EEFs was typically types of haplogroup G2a, and to a lesser extent H, T, J, C1a2 and E1b1, while their
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
was diverse. In the Balkans, the EEFs appear to have divided into two wings, who expanded further west into Europe along the
Danube The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
(
Linear Pottery culture The Linear Pottery culture (LBK) is a major archaeological horizon of the European Neolithic period, flourishing . Derived from the German ''Linearbandkeramik'', it is also known as the Linear Band Ware, Linear Ware, Linear Ceramics or Inc ...
) or the western
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
(
Cardial Ware Cardium pottery or Cardial ware is a Neolithic decorative style that gets its name from the imprinting of the clay with the heart-shaped shell of the '' Corculum cardissa '', a member of the cockle family Cardiidae. These forms of pottery a ...
). Large parts of Northern Europe and
Eastern Europe Eastern Europe is a subregion of the European continent. As a largely ambiguous term, it has a wide range of geopolitical, geographical, ethnic, cultural, and socio-economic connotations. The vast majority of the region is covered by Russia, whic ...
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. During the Chalcolithic and early
Bronze Age The Bronze Age is a historic period, lasting approximately from 3300 BC to 1200 BC, characterized by the use of bronze, the presence of writing in some areas, and other early features of urban civilization. The Bronze Age is the second prin ...
, the EEF-derived cultures of Europe were overwhelmed by successive invasions of
Western Steppe Herders 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 Eneolithic steppe around the turn of the 5th millennium BCE, ...
(WSHs) from the
Pontic–Caspian steppe The Pontic–Caspian steppe, formed by the Caspian steppe and the Pontic steppe, is the steppeland stretching from the northern shores of the Black Sea (the Pontus Euxinus of antiquity) to the northern area around the Caspian Sea. It extend ...
, who carried about 60%
Eastern Hunter-Gatherer In archaeogenetics, the term Eastern Hunter-Gatherer (EHG), sometimes East European Hunter-Gatherer, or Eastern European Hunter-Gatherer is the name given to a distinct ancestral component that represents descent from Mesolithic hunter-gatherers ...
(EHG) and 40% Caucasus Hunter-Gatherer (CHG) admixture. These invasions led to EEF
paternal A father is the male parent of a child. Besides the paternal bonds of a father to his children, the father may have a parental, legal, and social relationship with the child that carries with it certain rights and obligations. An adoptive fathe ...
DNA lineages in Europe being almost entirely replaced with EHG/WSH paternal DNA (mainly R1b and R1a). EEF
maternal ] A mother is the female parent of a child. A woman may be considered a mother by virtue of having given birth, by raising a child who may or may not be her biological offspring, or by supplying her ovum for fertilisation in the case of gestati ...
DNA (mainly haplogroup N) also heavily declined, being supplanted by steppe lineages, suggesting the migrations involved both males and females from the steppe. EEF mtDNA however remained frequent, suggesting admixture between WSH males and EEF females. EEF ancestry remains widespread throughout Europe, ranging from about 60% near the
Mediterranean Sea The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
(with a peak of 65% in the island of
Sardinia Sardinia ( ; it, Sardegna, label=Italian language, Italian, Corsican language, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese dialect, Algherese and Catalan languag ...
) and diminishing northwards to about 10% in northern Scandinavia.


Physical appearance

European hunter-gatherers were much
taller Taller is a Communes of France, commune in the Landes (department), Landes Departments of France, department in Nouvelle-Aquitaine in southwestern France. William II Sánchez of Gascony perpetrated a major defeat of the Vikings at Taller in 982 ...
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. The Early European Farmers are believed to have been mostly dark haired and dark eyed, and light skinned.


Studies


Ancient human genomes suggest three ancestral populations for present-day Europeans

identified Early European Farmers (EEFs) as a distinct ancestral component in a study published in ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are p ...
'' in 2014. Along with Ancient North Eurasians (ANEs) and
Western Hunter-Gatherer In archaeogenetics, the term Western Hunter-Gatherer (WHG), West European Hunter-Gatherer or Western European Hunter-Gatherer names a distinct ancestral component of modern Europeans, representing descent from a population of Mesolithic hunter-g ...
s, EEFs were determined to be one of the three major ancestral populations of modern-Europeans. About 44% of EEF ancestry was determined to come from a "
Basal Eurasian Basal Eurasian is a hypothetical lineage of anatomically modern humans with reduced, or zero, archaic hominin (Neanderthal) admixture compared to other ancient non-Africans. Basal Eurasians may originate from the Middle East or North Africa, and a ...
" population that split prior to the diversification of other non-African lineages.
Ötzi Ötzi, also called the Iceman, is the natural mummy of a man who lived some time between 3350 and 3105 BC, discovered in September 1991 in the Ötztal Alps (hence the nickname "Ötzi") on the border between Austria and Italy. Ötzi is believed to ...
was identified as EEF. EEFs were determined to be largely of
Near East The ''Near East''; he, המזרח הקרוב; arc, ܕܢܚܐ ܩܪܒ; fa, خاور نزدیک, Xāvar-e nazdik; tr, Yakın Doğu is a geographical term which roughly encompasses a transcontinental region in Western Asia, that was once the hist ...
ern origin, with slight WHG admixture. It was through their EEF ancestors that most modern Southern Europeans acquired their WHG ancestry. EEF ancestry in modern Europe ranged from 30% in the
Baltic States The Baltic states, et, Balti riigid or the Baltic countries is a geopolitical term, which currently is used to group three countries: Estonia, Latvia, and Lithuania. All three countries are members of NATO, the European Union, the Eurozone, ...
to 90% near the
Mediterranean Sea The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
.


Ancient mitochondrial DNA from the northern fringe of the Neolithic farming expansion in Europe sheds light on the dispersion process

found that the people of the
Funnelbeaker culture The Funnel(-neck-)beaker culture, in short TRB or TBK (german: Trichter(-rand-)becherkultur, nl, Trechterbekercultuur; da, Tragtbægerkultur; ) was an archaeological culture in north-central Europe. It developed as a technological merger of lo ...
of southern
Scandinavia Scandinavia; Sámi languages: /. ( ) is a subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, Norway, and Swed ...
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.


Massive migration from the steppe was a source for Indo-European languages in Europe

found that the amount of WHG ancestry among EEFs had significantly increased during the Middle Neolithic, documenting a WHG resurgence. It was found that EEF
Y-DNA The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abs ...
was typically types haplogroup G2a, while their
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
was diverse. During the Late Neolithic and
Early Bronze Age The Bronze Age is a historic period, lasting approximately from 3300 BC to 1200 BC, characterized by the use of bronze, the presence of writing in some areas, and other early features of urban civilization. The Bronze Age is the second prin ...
, G2a nearly disappears from Europe and is replaced with types of R1b and R1a, indicating a massive migration of people out of the
Pontic–Caspian steppe The Pontic–Caspian steppe, formed by the Caspian steppe and the Pontic steppe, is the steppeland stretching from the northern shores of the Black Sea (the Pontus Euxinus of antiquity) to the northern area around the Caspian Sea. It extend ...
. It has been suggested that this migration might be connected to the spread of
Indo-European languages The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, Du ...
in Europe.


A common genetic origin for early farmers from Mediterranean Cardial and Central European LBK cultures

found that the people of the
Linear Pottery culture The Linear Pottery culture (LBK) is a major archaeological horizon of the European Neolithic period, flourishing . Derived from the German ''Linearbandkeramik'', it is also known as the Linear Band Ware, Linear Ware, Linear Ceramics or Inc ...
(LBK) in Central Europe and people of the
Cardial Ware Cardium pottery or Cardial ware is a Neolithic decorative style that gets its name from the imprinting of the clay with the heart-shaped shell of the '' Corculum cardissa '', a member of the cockle family Cardiidae. These forms of pottery a ...
culture along the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
coast were descended from a homogenous community of EEFs with a common origin in the Balkans. EEF ancestors of the LBK people were expected to have migrated into Central Europe along the
Danube The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
river, while EEF ancestors of the Cardials were expected to have migrated along the Mediterranean coast. The Cardials appeared to have acquired a significant amount of hunter-gatherer ancestry during this process. Among modern populations,
Sardinians The Sardinians, or Sards ( sc, Sardos or ; Italian and Sassarese: ''Sardi''; Gallurese: ''Saldi''), are a Romance language-speaking ethnic group native to Sardinia, from which the western Mediterranean island and autonomous region of Italy de ...
and
Basque people The Basques ( or ; eu, euskaldunak ; es, vascos ; french: basques ) are a Southwestern European ethnic group, characterised by the Basque language, a common culture and shared genetic ancestry to the ancient Vascones and Aquitanians. Ba ...
were found to harbor the largest amount of EEF ancestry, which they probably acquired through descent from the Cardials.


Upper Palaeolithic genomes reveal deep roots of modern Eurasians

found that the ancestors of the EEF had split off from WHG around 43,000 BC, possibly through a migration of WHG into Europe. Around 23,000 BC, EEFs ancestors had again split into EEFs and Caucasian Hunter-Gatherers (CHGs). CHG admixture has been detected among people of the Yamnaya culture, who expanded massively throughout Europe from around 3,000 BC.


Genome-wide patterns of selection in 230 ancient Eurasians

found EEFs to be closely genetically related to
Neolithic The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several part ...
farmers of Anatolia. EEFs were found to have 7–11% more WHG ancestry than their Anatolian ancestors. This suggested that the EEFs belonged to a common ancestral population before their expansion into Europe. With regards
Y-DNA The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abs ...
, EEF males typically carried types of G2a. The study found that most modern Europeans can be modeled as a mixture of WHGs, EEFs and descendants of the
Yamnaya culture The Yamnaya culture or the Yamna culture (russian: Ямная культура, ua, Ямна культура lit. 'culture of pits'), also known as the Pit Grave culture or Ochre Grave culture, was a late Copper Age to early Bronze Age arch ...
. The Anatolian ancestors of the EEFs were found to be genetically different from modern peoples of the
Near East The ''Near East''; he, המזרח הקרוב; arc, ܕܢܚܐ ܩܪܒ; fa, خاور نزدیک, Xāvar-e nazdik; tr, Yakın Doğu is a geographical term which roughly encompasses a transcontinental region in Western Asia, that was once the hist ...
, and were instead shifted towards Europe. Middle Neolithic and Chalcolithic peoples of Iberia were found to be genetically similar to each other, and harbored reduced levels of EEF and increased levels of WHG ancestry compared to Early Neolithic individuals of the region. Peoples of the
Srubnaya culture The Srubnaya culture (russian: Срубная культура, Srubnaya kul'tura, ua, Зрубна культура, Zrubna kul'tura), also known as Timber-grave culture, was a Late Bronze Age 1850–1450 BC cultureParpola, Asko, (2012)"Format ...
and the earlier
Sintashta culture The Sintashta culture (russian: Синташтинская культура, Sintashtinskaya kul'tura), around 2050–1900 BCE, is the first phase of the Sintashta–Petrovka culture. or Sintashta–Arkaim culture,. and is a late Middle Bronze Ag ...
were found to harbor c. 15% EEF ancestry, suggesting that these cultures emerged through the eastward migration of Central European peoples with steppe-related ancestry.


The neolithic transition in the Baltic was not driven by admixture with early European farmers and extensive farming in Estonia started through a sex-biased migration from the steppe

found no evidence of EEF admixture among Neolithic populations of the eastern Baltic and the
East European forest steppe The East European forest steppe ecoregion (WWF ID: PA0419) is a patchwork of broadleaf forest stands and grasslands (steppe) that stretches 2,100 km across eastern Europe from the Ural Mountains in Ural, through Povolzhye, Central Russia to the ...
, suggesting that the hunter-gatherers of these regions avoided genetic replacement while adopting Neolithic cultural traditions. found that the people of the subsequent Corded Ware culture in the eastern Baltic carried steppe and hunter-gatherer-related paternal and autosomal ancestry, and some EEF maternal ancestry.


Ancient X chromosomes reveal contrasting sex bias in Neolithic and Bronze Age Eurasian migrations

found no significant evidence sex-bias in the admixure between EEFs and hunter-gatherers during the initial EEF expansion into Europe, although a larger number of hunter-gatherer females may have been incorporated into EEF communities during this phase. During Late Neolithic and Bronze Age however, a dramatic sex-bias was detected, suggesting heavy mixing between migrating males with steppe-related ancestry and local females with EEF ancestry.


Parallel palaeogenomic transects reveal complex genetic history of early European farmers

examined the genetic history of EEFs. It was found that the initial westward spread of the EEFs from the Balkans was accompanied only by slight admixture with hunter-gatherer populations. Peoples of Middle Neolithic and Chalcolithic Iberia were found to carry about 75% EEF ancestry and 25% WHG ancestry, more WHG ancestry than Early Neolithic Iberians. Significant reductions in EEF ancestry during the later phases of the Neolithic was also observed in Central Europe, particularly in the northern and eastern parts of the region.


The genomic history of Southeastern Europe

found that the EEFs had initially spread agriculture throughout Europe largely without admixture with local WHGs. It was proposed that this process had started through a single massive migration from Anatolia into the Balkans in the 7th millennium BC. The EEFs had subsequently split into two wings, one which spread northwards along the
Danube The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
through the
Linear Pottery culture The Linear Pottery culture (LBK) is a major archaeological horizon of the European Neolithic period, flourishing . Derived from the German ''Linearbandkeramik'', it is also known as the Linear Band Ware, Linear Ware, Linear Ceramics or Inc ...
, and another which spread westward across the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
coast through the
Cardial Ware Cardium pottery or Cardial ware is a Neolithic decorative style that gets its name from the imprinting of the clay with the heart-shaped shell of the '' Corculum cardissa '', a member of the cockle family Cardiidae. These forms of pottery a ...
culture. By 5600 BC, these cultures had brought agriculture to
Iberia The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, def ...
and
Central Europe Central Europe is an area of Europe between Western Europe and Eastern Europe, based on a common historical, social and cultural identity. The Thirty Years' War (1618–1648) between Catholicism and Protestantism significantly shaped the ar ...
. It was found that there was a significant increase hunter-gatherer ancestry in Iberia, Central Europe and the Balkans during the Middle Neolithic.. "We provide the first evidence for sex-biased admixture between hunter-gatherers and farmers in Europe, showing that the Middle Neolithic "resurgence" of hunter-gatherer-related ancestry in central Europe and Iberia was driven more by males than by females." While the slight mixture between EEFs and hunter-gatherers in the Early Neolithic appeared to have happened without sex-bias, increases in hunter-gatherer ancestry during the Middle Neolithic appeared to be largely the result of males with hunter-gatherer ancestry mixing with females with EEF ancestry. This conclusion was derived from the fact that examined Middle Neolithic Europeans overwhelmingly carried hunter-gatherer paternal lineages and EEF maternal lineages. Hunter-gatherer ancestry was even higher among Late Neolithic samples from the Cucuteni–Trypillia culture,
Funnelbeaker culture The Funnel(-neck-)beaker culture, in short TRB or TBK (german: Trichter(-rand-)becherkultur, nl, Trechterbekercultuur; da, Tragtbægerkultur; ) was an archaeological culture in north-central Europe. It developed as a technological merger of lo ...
and
Globular Amphora culture The Globular Amphora culture (GAC, (KAK); ), 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 ...
, which carried about 75-80% EEF ancestry while being dominated by hunter-gatherer paternal lineages. In the southern Balkans, the Middle Neolithic farmers display reduced levels of EEF ancestry increased amounts of ancestry related to Caucasian Hunter-Gatherers (CHGs), suggesting further gene flow from Anatolia, which continued into the
Bronze Age The Bronze Age is a historic period, lasting approximately from 3300 BC to 1200 BC, characterized by the use of bronze, the presence of writing in some areas, and other early features of urban civilization. The Bronze Age is the second prin ...
.


Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe

estimated that examined individuals at the Late Neolithic site of Kelif el Boroud,
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
, dated c. 3000 BC, carried about 50% EEF ancestry and 50% North African ancestry, were genetically predisposed to have light skin and light eyes, and entirely carried paternal and maternal lineages associated with EEFs. It was suggested that EEF ancestry had entered North Africa through Cardial Ware colonists from Iberia sometime between 5000 and 3000 BC. The examined samples of Kelif el Boroud were found to be closely related to
Guanches The Guanches were the indigenous inhabitants of the Canary Islands in the Atlantic Ocean some west of Africa. It is believed that they may have arrived on the archipelago some time in the first millennium BCE. The Guanches were the only nativ ...
of the Canary Islands. Additional amounts of EEF ancestry may have been brought to North Africa by the
Bell Beaker culture The Bell Beaker culture, also known as the Bell Beaker complex or Bell Beaker phenomenon, is an archaeological culture named after the inverted-bell beaker drinking vessel used at the very beginning of the European Bronze Age. Arising from ar ...
. The authors of the study suggested that the
Berbers , image = File:Berber_flag.svg , caption = The Berber ethnic flag , population = 36 million , region1 = Morocco , pop1 = 14 million to 18 million , region2 = Algeria , pop2 ...
of Morocco carried a substantial amount of EEF ancestry before the establishment of Roman colonies in Berber Africa.


Mitochondrial genomes reveal an east to west cline of steppe ancestry in Corded Ware populations

found that while females with steppe-related ancestry contributed to the formation of the Corded Ware culture in the eastern Baltic, the maternal lineages of Corded Ware culture on its western fringes were largely of EEF origin, suggesting that mixing that the westward expansion of the Corded Ware culture was characterized by the mixing of males with steppe-related ancestry and women with EEF ancestry.


The genomic history of the Iberian Peninsula over the past 8000 years

examined the
genetic history of the Iberian Peninsula The ancestry of modern Iberians (comprising the Spanish and Portuguese) is consistent with the geographical situation of the Iberian Peninsula in the south-west corner of Europe. As is the case for most of the rest of Southern Europe, the prin ...
. It was found that the peoples of Early Neolithic Iberia were largely of EEF ancestry. Peoples of the Middle Neolithic and Copper Age were found to harbor increased levels of WHG ancestry as compared to the Early Neolithic. Hunter-gatherer admixture was found to be higher in northern and central Iberia. Olalde argues that during the Bronze Age Iberia experienced a significant genetic turnover, with 100% of the paternal ancestry and 40% of the overall ancestry being replaced by peoples with steppe-related ancestry.


Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society

examined the remains of 24 individuals buried in
megalith A megalith is a large stone that has been used to construct a prehistoric structure or monument, either alone or together with other stones. There are over 35,000 in Europe alone, located widely from Sweden to the Mediterranean sea. The ...
s in northern and western Europe during the Middle Neolithic. They were found to be largely of EEF ancestry, although with significant amount of hunter-gatherer admixture, which appeared to be male-derived. The 17 samples of
Y-DNA The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abs ...
extracted belonged exclusively to the paternal haplogroup I, particularly I2, which are lineages associated with European hunter-gatherers. The evidence suggested that these societies were strongly
patrilineal Patrilineality, also known as the male line, the spear side or agnatic kinship, is a common kinship system in which an individual's family membership derives from and is recorded through their father's lineage. It generally involves the inheritanc ...
and socially stratified.


Ancient genomes indicate population replacement in Early Neolithic Britain

found that the farmers of the
Neolithic British Isles The Neolithic British Isles refers to the period of British, Irish and Manx history that spanned 4000 to 2,500 BCE. The final part of the Stone Age in the British Isles, it was a part of the greater Neolithic, or "New Stone Age", across E ...
had entered the region through a mass migration c. 4000 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.


Genetic history from the Middle Neolithic to present on the Mediterranean island of Sardinia

found that people of the Bronze Age
Nuragic civilization The Nuragic civilization, also known as the Nuragic culture, was a civilization or culture on Sardinia (Italy), the second largest island in the Mediterranean Sea, which lasted from the 18th century BC (Middle Bronze Age) (or from t ...
of
Sardinia Sardinia ( ; it, Sardegna, label=Italian language, Italian, Corsican language, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese dialect, Algherese and Catalan languag ...
carried about 80% EEF ancestry and 20% WHG ancestry. Strong evidence of genetic continuity was detected between Neolithic Sardinians and Bronze Age Sardinians.


A massacre of early Neolithic farmers in the high Pyrenees at Els Trocs, Spain

examined the remains of 9 farmers who had been systematically massacred at the site of Els Trocs, Spain c. 5300 BC. They were found to be genetically different from contemporary populations of Iberia, and were instead more similar to EEFs of Central Europe. The authors of the study suggested that they were migrant farmers from Central Europe who had been a victims of an ethnic cleansing carried out by local hunter-gatherer groups, or by other (either local or also migrant) farmer groups seeking to defend their territories.


Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers

found that Neolithic farmers in Western Europe had higher amounts of WHG ancestry than their Central European contemporaries. Neolithic farmers of France and Iberia were found to be closely related to contemporary farmers of the
British Isles The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isles, ...
, with whom they shared a relatively large amount of WHG ancestry. Examined farmers of Early Neolithic southern France exclusively carried the hunter-gatherer-derived paternal haplogroup I2, while the maternal lineages were mainly of EEF origin. Levels of Hunter-gatherer admixture among early farmers of France increased further during Middle Neolithic, reaching as high as 30% at some sites entirely dominated by hunter-gatherer paternal lineages. It was suggested that the increase was a result of migrations towards the northeast by farmers with elevated levels of hunter-gatherer ancestry.


Ancient genomes from present-day France unveil 7,000 years of its demographic history

found that earliest farmers of modern-day France were genetically similar to the Central European agriculturalists of the Linear Pottery Culture. It was found that the observed resurgence of WHG ancestry among European farmers in the Middle Neolithic happened very early and was relatively large in modern-day France.


A dynastic elite in monumental Neolithic society

examined a large number of individuals buried in
Neolithic Ireland The prehistory of Ireland has been pieced together from archaeological evidence, which has grown at an increasing rate over the last decades. It begins with the first evidence of permanent human residence in Ireland around 10,500 BC (although ...
. They 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. Peoples of the Irish and British Neolithic carried almost entirely the paternal haplogroup I-M284 (a WHG Y-haplogroup), suggesting that these societies were strongly
patrilineal Patrilineality, also known as the male line, the spear side or agnatic kinship, is a common kinship system in which an individual's family membership derives from and is recorded through their father's lineage. It generally involves the inheritanc ...
. A Neolithic royal buried at
Newgrange Newgrange ( ga, Sí an Bhrú) is a prehistoric monument in County Meath in Ireland, located on a rise overlooking the River Boyne, west of Drogheda. It is an exceptionally grand passage tomb built during the Neolithic Period, around 3200 BC, ...
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".


See also

*
Neolithic Europe The European Neolithic is the period when Neolithic (New Stone Age) technology was present in Europe, roughly between 7000 BCE (the approximate time of the first farming societies in Greece) and c.2000–1700 BCE (the beginning of the Bronze Age ...
* Neolithic decline *
Anatolian hunter-gatherers Epipaleolithic that inhabited central around 13,642-13,073 BCE. This population constitutes one of the three currently known ancestral genetic contributions of the present-day Europeans: Western Hunter Gatherers (WHG), Anatolian Neolithic Farmers ...


References


Bibliography

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Further reading

* * * * * * * * * * {{refend EEF Genetic history of Europe Neolithic Europe Modern human genetic history