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File:Genomic structure in Europeans dating back at least 36,200 years.jpg
Flowchart of Eurasian populations and their ancestries.[1]
File:Cro-Magnon man reconstruction.jpg
Reconstruction of a Cro-Magnon man. Upper Paleolithic and Mesolithic European early modern humans represent the first distinct West Eurasian lineage in Europe genetically different from Anatomically modern Homo sapiens remains found before such as the Ust'-Ishim man or the Goyet Cave remains.

In archaeogenetics, archaeology, and population genetics West Eurasian refers to the populations of Europe, the Middle East and other populations related to them.

West Eurasians owe the majority of their ancestry to a single largely verifiable migration out of Africa that occurred between 65,000 to 50,000 years BP, the same migratory episode believed to be ancestral to other modern non-African populations such as those of East Asia, Southeast Asia, and Siberia (known formerly as Mongoloid and in scientific bibliography as East Eurasian) as well as those of Oceania, and later than even earlier migrations that may not have been substantially ancestral to modern populations outside of Africa.[2][3][4] After this migratory event they quickly populated Europe, the Levant and Western Siberia defining Lithic stages such as the Ahmarian, Aurignacian, Levantine Aurignacian, Gravettian, Solutrean, Kebaran and Epigravettian cultures across the mentioned areas. Unifying characteristics of all West Eurasian lithic cultures could be Venus figurines, as well as similar technologies and tools. The first West Eurasian remains are related to the Cro-Magnon and similar Aurignacian remains found throughout Europe and the Levant.

West Eurasians share a higher degree of genetic similarity and consistency compared to other Non-Africans. However, they incorporated ancestry from several other human populations. Today there is no population on Earth that is purely West Eurasian descended.[citation needed] West Eurasians quickly divided into several related archaeogenetic groups such as Western Hunter-Gatherers, Ancient North Eurasians and Dzudzuana and Anatolian hunter-gatherers. These groups interacted with each other, usually violently and territorially but also amicably, generating further descendant groups such as Caucasian Hunter-Gatherers, Eastern Hunter-Gatherers, Iranian Hunter-Gatherers, Scandinavian Hunter-Gatherers as well as many others. At the advent of agriculture during the Neolithic Revolution, West Eurasian hunter-gatherer societies in Southern Europe, the Levant, the Caucasus, Siberia and Iranian Plateau became largely transhumant or sedentarian. Natufians or a population genetically related to natufians seems to have been the first to develop agriculture and to spread it through West Eurasia simultaneously to the dissemination of Afroasiatic languages through North Africa, the Levant, Arabia, Anatolia and the Caucasus. This expansion of new technologies caused the development of additional populations as hunter-gatherers integrated into the agricultural lifestyle such as Early European Farmers, Iranian Neolithic farmers, and Western Steppe Herders like the Yamnaya culture that spread Indo-European languages across West Eurasia; as well as Funnelbeakers, Globular Amphora culture, among others.[citation needed]

Archaeogenetic populations overwhelmingly descended from West Eurasians

Western Hunter-Gatherers (WHG)

File:1440x810 story-6e1e0e5e-5ae3-5a5e-b987-ced4efb13778 788983.jpg
Cheddar Man reconstruction.

Simultaneously to the introduction of West Eurasians into Europe a population not related to West Eurasians was already decaying demographically, this population is related to the remains found in the Goyet Caves and to the Mousterian and Châtelperronian lithic cultures. There is evidence of West Eurasians incorparting these decaying populations into theirs, such as the existence of Haplogroup C-V20 among modern Europeans and several Paleolithic remains such as the El Mirón Cave remains exhibit considerable amounts of non West Eurasian Ancestry. In archaeogenetics, the terms "Western Hunter-Gatherer", "West European Hunter-Gatherer" or "Western European Hunter-Gatherer", are the names given to a distinct ancestral component that represents descent from Mesolithic hunter-gatherers of Western, Southern and Central Europe. The term is often abbreviated as WHG. During the Mesolithic, the WHGs inhabited an area stretching from the British Isles in the west to the Carpathians in the east.

Ancient North Eurasians (ANE)

File:Sunghir Man.png
ANE derive most of their ancestry from a population related to the Sungir remains.

Ancient North Eurasians derive between 72 to 86% (78% according to Lazaridis[5]) of their ancestry from a particular West Eurasian population related to the remains found on the Sungir, Kostenki and Dolní Věstonice remains, all belonging to the Gravettian Period. The rest of ANE ancestry is East Eurasian. ANE were likely the first human population to exhibit Blonde hair.[6][7]

Dzudzuana and Anatolian hunter-gatherers (AHG)

Dzudzuana and AHG are two lineages from remains found in Ancient Anatolia and Prehistoric Georgia. Both lineages seem to be genetically related and derive most of their ancestry (72% according to Lazaridis) from West Eurasians, albeit incorporating genetic information from Basal Eurasians.[8]

Derived groups

Caucasian Hunter-Gatherer (CHG)

(CHG) is represented by the Satsurblia individual dated ~13 kya (from the Satsurblia cave in Georgia), and carried 36% ANE-derived admixture.[9] While the rest of their ancestry is derived from the Dzudzuana cave individual dated ~26 kya, which lacked ANE-admixture,[9] Dzudzuana affinity in the Caucasus decreased with the arrival of ANE at ~13 kya Satsurblia.[9]

Iran Neolithic (Iran_N)

(Iran_N) individuals dated ~8.5 kya carried 50% ANE-derived admixture and 50% Dzudzuana-related admixture,[9] marking them as different from other Near-Eastern and Anatolian Neolithics who didn't have ANE admixture.[9] Iran Neolithics were later replaced by Iran Chalcolithics, who were a mixture of Iran Neolithic and Near Eastern Levant Neolithic.[10]

Eastern Hunter-Gatherer (EHG)

(EHG) is a lineage derived predominantly (75%) from ANE and 25% from WHG[10] It is represented by two individuals from Karelia, one of Y-haplogroup R1a-M417, dated c. 8.4 kya, the other of Y-haplogroup J, dated c. 7.2 kya; and one individual from Samara, of Y-haplogroup R1b-P297, dated c. 7.6 kya. This lineage is closely related to the ANE sample from Afontova Gora, dated c. 18 kya. After the end of the Last Glacial Maximum, the Western Hunter-Gatherers (WHG) and EHG lineages merged in Eastern Europe, accounting for early presence of ANE-derived ancestry in Mesolithic Europe. Evidence suggests that as Ancient North Eurasians migrated West from Eastern Siberia, they absorbed Western Hunter-Gatherers and other West Eurasian populations as well.[7]

Scandinavian Hunter-Gatherer (SHG)

File:Scandinavian Hunter-Gatherer.jpg
Reconstruction of an Scandinavian Hunter-Gatherer woman.

(SHG) is the name given to a distinct ancestral component that represents descent from Mesolithic hunter-gatherers of Scandinavia.[a][12] 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.[citation needed]

Early European Farmers/Early Neolithic Farmers/European Neolithic Farmers (EEF/ENF)

File:Ötzi EEF.jpg
Reconstruction of Ötzi, a mummy adscribed to the ENF linage. Due to their diet and lifestyle, EEF are among the first populations to experience a degree of depigmentation similar to that of modern Europeans.

EEF and ENF are diffuse terminologies utilized to denote several remains and populations related to each other, usually as a result of interactions between Western Hunter-Gatherers (WHG) and Dzuduana/Anatolian hunter-gatherers (AHG). Due to several contemporary migrations and invasions, the EEF/ENF genetic component varies greatly, with some studies identifying EEF as being up to 44% Basal Eurasian,[13] while other studies show EEF being almost genetically indistinguishable from AHG (72% WHG and 28% Basal Eurasian), and some other estudies showing EEF/ENF being largely affected by WHG and EHG migrations into Arabia through the Caucasus that replaced most of the male genetic lineage for the European derived[citation needed] Haplogroup I[disambiguation needed] and Haplogroup J-M172, and thus, very related to Epigravettian remains such as the Ripari Villabruna complex.[14]

A genetic study published in Nature in November 2015 found EEFs to be closely genetically related to Neolithic farmers of Anatolia. EEFs were found to have 7–11% more WHG ancestry than their Anatolian relatives. This suggested that the EEFs belonged to a common ancestral population before their expansion into Europe. With regards Y-DNA, EEF males typically carried types of G2a. The study found that most Europeans could be modeled as a mixture of WHGs, EEFs and descendants of the Yamnaya culture.[b]

Western Steppe Herders (WSH)

File:Yamnaya male (3) (2).jpg
Forensic facial reconstruction of a male from the Yamnaya culture. Western Steppe Herder (WSH) ancestry is believed to have been spread throughout large parts of Eurasia by the Yamnaya.[16]

WSHs are considered descended from Eastern Hunter-Gatherers (EHGs) who received some admixture from Caucasian Hunter-Gatherers (CHGs) during the Neolithic. The Y-DNA of the WSHs was mostly types of R1a and R1b, which are EHG lineages, suggesting that CHG admixture among the WSHs came through EHG males mixing with CHG females.[17] Around 3,000 BC, people of the Yamnaya culture, who belonged to the WSH cluster, embarked on a massive expansion throughout Eurasia, which might have resulted in the dispersal of Indo-European languages. WSH ancestry from this period is often referred to as Steppe Early and Middle Bronze Age (Steppe EMBA) ancestry.[c]

Archaeogenetic populations partially descended from West Eurasians

Ancient Beringian/Ancestral Native American (AB/ANA)

AB and ANA are specific archaeogenetic lineages, based on the genome of an infant found at the Upward Sun River site (dubbed USR1), dated to 11,500 years ago.[18] Between 14% and 38% of Native American ancestry may originate from gene flow from the Mal'ta Buret people (Ancestral North Eurasians). This variation is caused by the penetration of posterior Siberian migrations into the Americas, with the lowest percentages of ANE ancestry found in Eskimos and Alaskan Natives, as these groups are the result of migrations into the Americas roughly 5,000 years ago.[19]

A more recent study in 2020 found the oldest relative of modern Native Americans in Siberia, a 11,000 year old individual that shared close to two thirds of his ancestry to modern Native Americans and genetically related to East Asians. The European-related ancestry, the Ancestral North Eurasian component, is lower than estimated before and was itself found to be the admixture of East Asian and European lineages.[20]

Ancestral South Indian/Ancient Ancestral South Indian (ASI/AASI)

Results of studies based upon autosomal DNA variation have also been varied. In a major study (2009) using over 500,000 biallelic autosomal markers, Reich hypothesized that the modern South Asian population was the result of admixture between two genetically divergent ancestral populations dating from the post-Holocene era. These two "reconstructed" ancient populations he termed "Ancestral South Indians" (ASI) and "Ancestral North Indians" (ANI). According to Reich: "ANI ancestry is significantly higher in Indo-European than Dravidian speakers, suggesting that the ancestral ASI may have spoken a Dravidian language before mixing with the ANI." While the ANI is genetically close to Middle Easterners, Central Asians and Europeans, the ASI is not closely related to groups outside of the subcontinent. As no "ASI" ancient DNA is available, the indigenous Andamanese Onge are used as an (imperfect) proxy of ASI, though nevertheless distinct from (according to Reich et al., the Andamanese, though distinct from them, are the closest living population to ASI). According to Reich et al.[citation needed], both ANI and ASI ancestry are found all over the subcontinent (in both northern and southern India) in varying proportions, and that “ANI ancestry ranges from 39-71% in India, and is higher in traditionally upper caste and Indo-European speakers."

Moorjani et al. 2013 state that the ASI, though not closely related to any living group, are "related (distantly) to indigenous Andaman Islanders." Moorjani et al. also suggest possible gene flow into the Andamanese from a population related to the ASI. The study concluded that “almost all groups speaking Indo-European or Dravidian languages lie along a gradient of varying relatedness to West Eurasians in PCA (referred to as “Indian cline”)”.[21]

A 2013 study using the single-nucleotide polymorphism (SNP), shows that the genome of Andamanese people (Onge) is closer to those of other Oceanic Negrito groups than to that of South Asians.[22]

According to Basu et al. 2016, further analysis revealed that the genomic structure of mainland Indian populations is best explained by contributions from four ancestral components. In addition to the ANI and ASI, Basu et. al (2016) identified two ancestral components in mainland India that are major for the Austro-Asiatic-speaking tribals and the Tibeto-Burman speakers, which they denoted as AAA (for “Ancestral Austro-Asiatic”) and ATB (for "Ancestral Tibeto-Burman") respectively. The study also infers that the populations of the Andaman Islands archipelago form a distinct ancestry, which "was found to be coancestral to Oceanic populations".[23]

The cline of admixture between the ANI and ASI lineages is dated to the period of c. 4.2–1.9 kya by Moorjani et al. (2013), corresponding to the Indian Bronze Age, and associated by the authors with the process of deurbanisation of the Indus Valley Civilization and the population shift to the Gangetic system in the incipient Indian Iron Age. Basu et al. (2003) suggests that "Dravidian tribals were possibly widespread throughout India before the arrival of the Indo-European-speaking nomads" and that "formation of populations by fission that resulted in founder and drift effects have left their imprints on the genetic structures of contemporary populations". The geneticist PP Majumder (2010) has recently argued that the findings of Reich et al. (2009) are in remarkable concordance with previous research using mtDNA and Y-DNA:[citation needed]

In general, Ancestral South Indians formed as mixture of "Indus Periphery-related" groups (Iranian agriculturalists) who moved south and mixed with AASI-related ancestry.[24]

Natufians

File:Out of Africa migratory model.jpg
Out of Africa Migratory model depicting Anatomically modern Homo sapiens dispersals. West Eurasians, in common with other modern non-African populations, descend from the final and most impactful major migratory movement of Homo sapiens from Africa, depicted here as happening 55,000 years ago, entering Europe 45,000 years ago coinciding with the start of the Aurignacian.[citation needed]

According to ancient DNA analyses conducted by Lazaridis et al. (2016) on Natufian skeletal remains from present-day northern Israel, the Natufians carried the Y-DNA (paternal) haplogroups E1b1b1b2(xE1b1b1b2a,E1b1b1b2b) (2/5; 40%), CT (2/5; 40%), and E1b1(xE1b1a1,E1b1b1b1) (1/5; 20%).[25][26] In terms of autosomal DNA, these Natufians carried around 50% of the Basal Eurasian (BE) and 50% of Western Eurasian Unknown Hunter Gather (UHG) components. However, they were slightly distinct from the northern Anatolian populations that contributed to the peopling of Europe, who had higher Western Hunter Gatherer (WHG) inferred ancestry. Natufians were strongly genetically differentiated[27] from Neolithic Iranian farmers from the Zagros Mountains, who were a mix of Basal Eurasians (up to 62%) and Ancient North Eurasians (ANE). This might suggest that different strains of Basal Eurasians contributed to Natufians and Zagros farmers,[28][29][30] as both Natufians and Zagros farmers descended from different populations of local hunter gatherers. Contact between Natufians, other Neolithic Levantines, Caucasus Hunter Gatherers (CHG), Anatolian and Iranian farmers is believed to have decreased genetic variability among later populations in the Middle East. The scientists suggest that the Levantine early farmers may have spread southward into East Africa, bringing along Western Eurasian and Basal Eurasian ancestral components separate from that which would arrive later in North Africa. According to their results, the Natufians shared no genetic affinity to present-day sub-Saharan Africans. However the scientists state that they were unable to test for affinity in the Natufians to early North African populations using present-day North Africans as a reference because present-day North Africans owe most of their ancestry to back-migration from Eurasia.[25][31]

Ancient DNA analysis has confirmed ancestral ties between the Natufian culture bearers and the makers of the Epipaleolithic Iberomaurusian culture of the Maghreb,[32] the Pre-Pottery Neolithic culture of the Levant,[32] the Early Neolithic Ifri n'Amr or Moussa culture of the Maghreb,[33] the Savanna Pastoral Neolithic culture of East Africa,[34] the Late Neolithic Kelif el Boroud culture of the Maghreb,[33] and the Ancient Egyptian culture of the Nile Valley,[35] with fossils associated with these early cultures all sharing a common genomic component.[33]

A 2018 analysis of autosomal DNA using modern populations as a reference, found the Natufians to have a predominant mixture of ancestral components from Western Eurasia and North Africa as well as a small (ca 6.8%) East African-related ancestry (showing affinities to the Omotic peoples of southern Ethiopia). It is suggested that this East African component may have been associated with the spread of Y-haplogroup E (particularly Y-haplogroup E-M215, also known as "E1b1b") lineages to Western Eurasia.[36]

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