Learning Materials
Introductuory Articles to Population Genetics
Out of Africa (OoA) and East-West Eurasian Split
It is widely acknowledged that the ancestors of all the non-African (Eurasian) populations migrated out of the African continent approximately 70-60 kya (Vallini et al. 2022). During their initial expansion into Eurasia, possibly somewhere in the Near East, they interbred with the Neanderthals, resulting in an estimated 2-3% Neanderthal admixture in the majority of the later populations (Posth et al. 2023). This initial group of Eurasian population further split into West and East Eurasian groups, believed to have occurred between 45 to 38 thousand years ago.
The oldest Homo Sapiens specimen sequenced to date is a female individual from Zlaty Kun (Czech Republic) is radiocarbon dated to over 45 ka. The Zlaty Kun individual represents an ancestral lineage that is basal to both West and East Eurasians (Vallini et al. 2022). In contrast, the second oldest human samples from the Bacho Kiro cave in Bulgaria, dated around 45 thousand years ago, exhibit a closer genetic affinity with ancient and contemporary East Asians than with Europeans. Moreover, individuals from Bacho Kiro display a notably higher proportion of Neanderthal admixture, suggesting recent interbreeding with Neanderthals within the last few generations (Vallini et al. 2022). On the other hand, the genetic positioning of the Ust’Ishim individual (45 ka) from Omsk Oblast, Russia, contemporaneous with Bacho Kiro, remains ambiguous. Nevertheless, statistical support leans towards the East Eurasian lineage for Ust’Ishim (Vallini et al. 2022, supplementary information).
So, to summarise, the latest findings on the oldest human samples indicate that the initial migration wave out of Africa, as represented by Zlaty Kun, had already populated Europe prior to 45 ka. Around 45 ka, individuals with a genetic affinity to East Eurasians emerged in locations such as Bacho Kiro cave in Bulgaria and Ust’Ishim in Russia. Subsequently, Oase1 in Romania (40 ka, likely descended from Bacho Kiro) and the Tianyuan individual in China (40 ka) also displayed similar genetic profiles.
It is also noteworthy that the ancestral lineage represented by individuals predating 40 ka (with the exception of Tianyuan) appears to have either become extinct or assimilated into later migrating populations (Posth et al. 2023).
Some time after the appearance of the East Eurasian related samples, we see the occurrence of the samples carrying ancestries that derive primarily from the lineage leading to present-day Europeans (West Eurasian). The oldest human genomes reflecting this lineage include Kostenki 14 (38 ka) from western Russia and GoyetQ-116-1 (35 ka) from Goyet cave in Belgium. The ancestral signature found in Kostenki 14 later contributed to the Vestonice cluster (named after the Dolní Věstonice site in Czechia), associated with the archaeologically defined Gravettian Culture (33-36 ka). This ancestral lineage was shared among individuals from the Gravettian Culture in Central and Southern Europe but disappeared after the Last Glacial Maximum (LGM).
Conversely, the ancestry related to GoyetQ-116-1 reappeared after the LGM in individuals associated with the archaeological Magdalenian Culture (19-14 ka) from Iberia across Central Europe to Eastern Europe. This lineage was later largely replaced by the Epigravettian-associated (24-12 ka) Villabruna (WHG) cluster (Posth et al. 2023).
- Vallini, L., et al. (2022). Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa. Genome Biology and Evolution, Volume 14, Issue 4, April 2022, evac045, https://doi.org/10.1093/gbe/evac045
- Posth, C., et al. (2023). Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers. Nature, 615(7950), 117–126. https://doi.org/10.1038/s41586-023-05726-0
Europe: From Paleolithic to Neolithic
In this section we will briefly go through the genetic breakpoints of Europe, beginning from the Initial Paleolithic Age to the Neolithic farmer migrations. The information below relies on two extensive recent studies: Posth et al. 2023 and Allentoft et al. 2024.
Overview
The genetic history of Paleolithic Europe can be roughly categorised into two periods: pre and post the Last Glacial Maximum (25,000-19,000 BP). In the pre-Last Glacial Maximum era, Europe had two distinct ancestral groups. The first group is represented by the individual Goyet-Q116-1 (35,000 BP) in Belgium, and the second group is represented by individuals from Kostenki 14 (32,000 BP) and Sunghir (34,000 BP) in western Russia.
The ancestry associated with Goyet-Q116-1 corresponds to the Aurignacian Period, a period that later transitioned into the Gravettian Culture (33-26,000 BP), giving rise to two ancestral clusters in Europe. Central, east, and southern Europe received a mixture related to Sunghir, resulting in an estimated 64% Sunghir and 36% Goyet-Q116-1-like admixture, known as the Vestonice Cluster. In contrast, the genetic cluster in west and southwest Europe, known as the Fournol Cluster, was closely linked to the ancestry represented by Goyet-Q116-1. This ancestry persisted during and after the LGM in west and southwest Europe.
The LGM period in southwest Europe is archaeologically associated with the Solutrean Culture. Samples from this period exhibit an intermediate genetic profile between preceding Fournol ancestry, associated with Gravettian Culture and succeeding Magdalenian Cultures, represented by the oldest Magadelanian associated sample El Miron.
Magdenalian Culture had two genetic clusters in itself. The first one is the Goyet-Q2 cluster with samples 15.000 years old (from France, Belgium, Germany) and the second one is the 19.000 years old El Miron cluster from Spain. Both clusters are distantly related to Goyet-Q116-1 although having significant contribution from the Eppi-Gravettian associated Villabruna Cluster.
After the LGM, the Eppi-Gravettian culture spread widely in south and southeast Europe. The ancestry related to this culture is called Villabruna Cluster, which, also will be known as the Western European Hunter-Gatherer (WHG) ancestry. This ancestry was not related to the preceding Gravettian associated Vestonice Cluster, which seemingly disappeared after the LGM. The Villabruna Cluster later largely replaced Magdelanian culture which was spread in southwest, west and central Europe. Villabruna/Oberkassel/WHG ancestry dominated most part of Europe until the Neolithic Age and then largely replaced by the Anatolian Neolithic Farmer migrations.
Timeline
Pre-LGM
The oldest genomes carrying ancestries that derive primarily from the lineage leading to present-day Europeans are Kostenki 14 (from 37 ka, with uncertain archaeological association from western Russia), Goyet Q116-1 (35 ka, Aurignacian-associated from Belgium) and Bacho Kiro 1653 (35 ka, probably Aurignacian-associated from Bulgaria)
Gravettian Period (33–26 ka):
Věstonice genetic cluster: Named after the Dolní Věstonice site in Czechia. Comprising Gravettian associated individuals from central-east and southern Europe (Dolní Věstonice, Pavlov, Krems-Wachtberg, Paglicci and Ostuni). Emerged by the genetic contribution from a Sunghir/Kostenki 14-related source. This genetic signature is shared among individuals associated with the archaeologically defined Gravettian culture in central and southern Europe and seemingly disappeared after the Last Glacial Maximum
Fournol genetic cluster: Comprising Gravettian-associated individuals from western and southwestern European sites (Ormesson, La Rochette, Fournol and two Serinyà cave sites (Mollet III and Reclau Viver).Fournol cluster is closely related to Aurignacian-associated individuals from Belgium dated to 35 ka (Goyet Q116-1 and Goyet Q376-3 individual).
During this period genetically intermediate samples between Vestonice and Fournol are also observed.
During LGM
Solutrean culture (24–19 ka): Emerged during the LGM in southwest Europe. The LGM is considered to have caused a demographic decline in large parts of Europe, with populations retracting to southern latitudes, to the Iberian peninsula and southern France.
Epigravettian culture: (24–12 ka): During the LGM Epigravettian culture seems to have gradually spread from the Balkans to Italy. The distinct ancestry represented by Epigravettian associated individuals is called Villabruna or Oberkassel cluster. This ancestry initially replaced the Vestonice Cluster in the region and later largely replaced Magdalenian associated ancestry in much of Europe.
Post LGM
Magdalenian Culture (19-14 ka): Succeeded from Solutrean Culture in Iberia and France and spread across eastern and central Europe. Ancestry represented by Magdalenian associated individuals (Goyet Q2) is distantly linked to the Goyet Q116-1 individual from Belgium dated to 35 ka. Magdalenian Culture and the ancestry associated with it largely replaced by the Epigravettian Culture
Epigravettian culture: (24–12 ka): Upon mostly replacing the Magdalenian Culture, Epigravettian associated Villabruna/Oberkassel (WHG) ancestry became the dominant and most widespread ancestry in Europe until the Mesolithic era. Around the Mesolithic, the ANE related population mixed with WHG appears in Eastern Europe, giving rise to Eastern European Hunter-Gatherers (EHG/Sidelkino Cluster).
Neolithic Transition
The arrival of Anatolian Farmer-related ancestry in different regions of Europe spans an extensive time period of over 3,000 years, from its earliest evidence in the Balkans (Lepenski Vir) at ∼8,700 BP to c. 5,900 BP in Denmark. These Neolithic migrations caused an East-West distinction along a boundary zone running from the Black Sea to the Baltic, causing the “Great Divide” in Europe. In the west of this division, Anatolian Farmer-related ancestry was largely replacing the local hunter gatherers (WHG).
However, on the eastern side of the ‘“Great Divide’” no ancestry shifts can be observed during this period. In the East Baltic region, the Ukraine and Western Russia, local HG ancestry (represented by the EHG) prevailed until ∼5,000 BP without noticeable input of Anatolian-related farmer ancestry.
- Allentoft, M.E., et al. (2024). Population genomics of post-glacial western Eurasia. Nature 625, 301–311. https://doi.org/10.1038/s41586-023-06865-0
- Posth, C. et al. (2023). Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers. Nature, 615(7950), 117–126. https://doi.org/10.1038/s41586-023-05726-0
Ancient North Eurasians (ANE)
The Ancient North Eurasian (ANE) is a distinctive ancestral component that emerged during the Upper Paleolithic Age. It is represented by a boy known as MA-1, associated with the Mal’ta-Buret’ Culture (24,000 BP) in Irkutsk Oblast, Russia, and individuals from Afontova Gora (18,000 BP) situated on the left bank of the Yenisei River in Krasnoyarsk, Russia. The ANE population is considered intermediate between West and East Eurasians and can be modeled as a combination of 50% Sunghir/Kostenki14 and 50% Tianyuan-like ancestry (Posth et al., 2023). This population played a pivotal role in shaping later ancient populations in Europe, the Middle East, and Siberia.
The ANE is believed to have descended either from Ancient North Siberians (ANS), represented by individuals associated with the preceding Yana Culture (32,000 BP) in Sakha, Russia, or from a common older population that is ancestral to both ANE and ANS. Yana individuals, much like those of the ANE, can be modeled as either 78% Sunghir and 22% Tianyuan or as a 50% Sunghir/Kostenki14 and 50% Tianyuan-like combination.
The ANE population’s one of the most significant roles was forming up the Eastern European Hunter Gatherers (EHG) by mixing partially with the WHG and Upper Paleolithic Caucasus Hunter Gatherer-related source in Eastern Europe, some time around the Mesolithic Age. The ANE ancestry was later exclusively carried by Western Siberian Hunter Gatherers (WSHG) primarly represented by an individual called Tyumen_HG, Botai Culture and Bronze Age Tarim populations (Tarim_EMBA) from Tarim Basin, China.
- Posth, C. et al. (2023). Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers. Nature, 615(7950), 117–126. https://doi.org/10.1038/s41586-023-05726-0
- Vallini, L. et al. (2022). Genetics and Material Culture Support Repeated Expansions into Paleolithic Eurasia from a Population Hub Out of Africa. Genome Biology and Evolution, Volume 14, Issue 4, April 2022, evac045, https://doi.org/10.1093/gbe/evac045
Basal Eurasians
Basal Eurasians are considered a “ghost” population believed to have significantly influenced ancient populations in the Middle East and North Africa during the Upper Paleolithic to Mesolithic periods. The earliest sequenced sample with Basal Eurasian ancestry (~24%), known as NEO283, dates back to the Upper Paleolithic era (26,000 BP) from Kotias Cave in Georgia, with the remaining ancestry attributed to West Eurasian Upper Paleolithic hunter-gatherers. (Allentoft et al., 2024)
Between the Upper Paleolithic and Mesolithic periods, there is an observed increase in Basal Eurasian ancestry in the Caucasus and Iran, coupled with an Ancient North Eurasian (ANE)-related admixture. This results in approximately 35% Basal Eurasian + ANE and 65% Basal Eurasian + ANE admixture in Mesolithic Caucasus Hunter Gatherers (CHG) and Mesolithic Iran (HotuIIIb) samples, respectively. However, the Basal Eurasian admixture decreases to less than 50% in Neolithic Iran (Lazaridis et al., 2016).
In contrast to the Caucasus and Iran, Basal Eurasian ancestry in Anatolian and Natufian Hunter Gatherers in the Levant is accompanied by Western Hunter-Gatherer (WHG)-like admixture. Anatolian Hunter Gatherers and their direct descendants, Anatolian Neolithic Farmers, are estimated to have approximately 25% Basal Eurasian + WHG ancestry (Feldman et al., 2019) while Natufian Hunter Gatherers exhibit around 45% Basal Eurasian + WHG-like ancestry. It remains unclear whether this WHG-like ancestry later migrated from South Europe to the Middle East or was already present in the region (Lazaridis et al., 2016, supplementary information).
Ferreira et al. (2021) proposed that Basal Eurasians and the ancestors of East and West Eurasians were genetically close, with Basal Eurasians likely descending from the same group that left Africa but split earlier, either in southwest Asia or in Africa before the Out-of-Africa (OoA) event. In the latter scenario, they followed the ‘southern route’ in the Arabian Peninsula and took refuge somewhere in the Persian gulf while the sister group followed ‘northern route’ mixed with the Neanderthals in northern Arabian Peninsula before moving further into Europe and Asia. Ferreira’s estimations also suggest that Iberomaurissians from the Tarofalt Culture (15,000 – 13,000 BP) in Morocco carry approximately 60% Basal Eurasian ancestry. Additionally, in contrast to Lazaridis et al. (2016), Natufians are estimated to have above 20% Basal Eurasian ancestry, while CHGs have approximately 40%.
- Allentoft, M.E., et al. (2024). Population genomics of post-glacial western Eurasia. Nature 625, 301–311. https://doi.org/10.1038/s41586-023-06865-0
- Ferreira, J. C., et al. (2021). Projecting Ancient Ancestry in Modern-Day Arabians and Iranians: A Key Role of the Past Exposed Arabo-Persian Gulf on Human Migrations. Genome biology and evolution, 13(9), evab194. https://doi.org/10.1093/gbe/evab194
- Feldman, M., et al. (2019). Late Pleistocene human genome suggests a local origin for the first farmers of central Anatolia. Nat Commun 10, 1218. https://doi.org/10.1038/s41467-019-09209-7
- Lazaridis, I., et al. (2016). Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419–424 https://doi.org/10.1038/nature19310
Asia: Primary Ancestral Groups
The genetic history of East Eurasian populations is a complex topic and still under an ongoing debate. Here, we will try to summarise the major ancestral clusters that played a significant role in shaping the later populations in the region under the guidance of the latest studies.
Abbreviations
M: Mesolithic
N: Neolithic
EN: Early Neolithic
EMN: Early-Middle Neolithic
MN: Middle Neolithic
LN: Late Neolithic
LNBA: Late Neolithic-Bronze Age
Northeast Asia
Khairygas_16.7kya
One of the oldest samples recovered from Northeast Asia, this sample may be a near-unadmixed representative of Ancient Paleosiberian (APS) ancestry and likely to be a near-unadmixed descendant of a population closely related to the founding population of the Americas. This sample can be modelled as ~97% Native American-like (represented by Peru_Laramate_900B) and ~7% ANE-related (represented by AfantovaGora3). (Zeng et al. 2023, sup. mat. p. 172-173)
Ust_Kyakhta_14kya
The second oldest sample in the APS cluster, this individual can be modelled as ~53-69% preceding Khairygas plus an undefined East Asian source, possibly related to KhatystyrCave_M_10.2kya. (Zeng et al. 2023, sup. mat. p. 173)
KhatystyrCave_M_10.2kya
Appearing to be of mixed ancestry, in a 3-way model, this individual primarily inherits the majority of its genetic heritage from China_AmurRiver_Mesolithic (~57-78% or alternatively as ~61% ancestry from China_AmurRiver_Mesolithic + ~24% ancestry from Cisbaikal_LNBA + ~15% from China_SEastAsia_Coastal_EN), with the remaining portion originating from Ust_Kyakhta and other East Asian source (either China_NEastAsia_Inland_EN or China_SEastAsia_Coastal_EN), (Zeng et al. 2023, sup. mat. p. 173-174).
Kolyma_M_10.1kya
While models for Kolyma_M_10.1kya are quite varied, they are highly consistent in suggesting that this population is mostly descended from Ancient Paleosiberians (~65-73% Khairygas_16.7kya-like) and is less admixed with East Asian ancestry compared to Ust_Kyakhta_14kya. The remaining East Asian ancestry of Kolyma_M_10.1kya seems to be derived from either Yumin or Amur related sources (Zeng et al. 2023, sup. mat. p. 174/196).
Dzhilinda1_M_N_8.4kya
This individual can be modeled as having all its ancestry coming from Ust_Kyakhta_14kya, or alternatively as an admixture of Kolyma_M_10.1kya or Khaiyrgas and an East-Asian-related source with much ancestry from KhatystyrCave_M_10.2kya or Transbaikal_EMN_old (Zeng et al. 2023, sup. mat. p. 176).
Transbaikal_EMN_old
Consisting of two individuals within the Transbaikal_EMN cluster (irk007.SG and cta016.SG), both older than the others at approximately 8,300 and 8,800 years before present (BP) respectively. This genetic makeup can be represented as a two-way admixture between the Yumin individual (China_NEastAsia_Inland_EN) and KhatystyrCave_M_10.2kya (Zeng et al. 2023, sup. mat. p. 176).
Altai_N_old
The oldest individuals under the Altai_N group label (uniting Kemerovo_N, Firsovo_N, and West_Siberia_N—extremely similar populations found in the Altai region) older than ~8000 BP (including individuals I2137 and I13098), can be successfully modeled as a 2-way admixture between a minor fraction of ancestry from AG3 and a major fraction of ancestry from the three APS populations Khaiyrgas, Ust_Kyakhta_14kya, Kolyma_M_10.1kya. The same models pass in set B; no simpler models pass (Zeng et al. 2023, sup. mat. p. 177).
Altai_N
This group, which comprises all individuals in the Altai_N group label (all individuals labeled Kemerovo_N, Firsovo_N, and West_Siberia_N) younger than 8000 BP, can be modeled as descending completely from the older individuals in this cluster (Altai_N_old), (Zeng et al. 2023, sup. mat. p. 178).
Transbaikal_EMN
Transbaikal_EMN, consisting of younger individuals in the Transbaikal_EMN cluster can be either modeled as descending completely in a one-way model from Transbaikal_EMN_old or two-way model with small additional contribution from China_NEastAsia_Inland_EN (represented by Yumin individual), (Zeng et al. 2023, sup. mat. p. 178).
China_AmurRiver_N
This population, consisting of individuals from the Amur River Basin region younger than ~8000 BP, can be modeled as descending almost entirely (~77-98%) from prior populations of the Amur River region (China_AmurRiver_EarlyN or China_AmurRiver_Mesolithic) in multiple 2-way models with the addition of a small contribution from one other, typically Northeast Asian or APS source (Zeng et al. 2023, sup. mat. p. 178).
The individuals labelled as Russia_Boisman_MN and Devils_Gate_N from the Amur Basin region are also associated with this ancestral cluster. Present day Tungusic populations from the region also show strong correlation with this ancestry.
Cisbaikal_EN
Models of Cisbaikal_EN, which includes individuals from the Cisbaikal region from ~8000BP to ~6800BP, with major contributions from Transbaikal_EMN_old (~65%-79%), and minor contributions from an APS population (Ust_Kyakhta_14kya ~35% or Khairygas_16.7kya ~31%), (Zeng et al. 2023, sup. mat. p. 178).
Mongolia_N_North
A two-way model for Mongolia_N_North, from the northern region of the Mongolian plateau just south of Lake Baikal approximately 7500 BP, as an admixture between Transbaikal_EMN_old (~65%) + China_NEastAsia_Inland_EN (~35%) an alternatively with the addition of a small fraction (<7%) of ancestry from an ANE-rich source (either Altai_N_old or from a APS-related population such as Khairygas_16.7kya, Ust_Kyakhta_14kya, Dzhilinda1_M_N_8.4kya or Kolyma_M_10.1kya), (Zeng et al. 2023, sup. mat. p. 178)
Syalakh-Belkachi
The Syalakh-Belkachi population in the Middle Lena river valley in Central Yakutia from 6800-6300 BP can be modeled as descending entirely from the Mesolithic Northern Buryatian Dzhilinda1_M_N_8.4kya. On the other passing 2-way models with higher p-values, where Syalakh-Belkachi continues to demonstrate majority descent from Dzhilinda1_M_N_8.4kya with a minor contribution (~10-20%) from a broadly East-Asian-related population whose identity is poorly resolved. In other passing models, major contribution from Dzhilinda1_M_N_8.4kya (~76%) and a minor contribution from a population rich in Inland Northeast Asian ancestry (represented by the Yumin individual under the group label China_NEastAsia_Inland_EN), either Cisbaikal_EN, Transbaikal_EMN_old or Mongolia_N_North (~24%) is observed (Zeng et al. 2023, sup. mat. p. 178-179)
This population shows high affinity with the Paleo Eskimo-Inuit cultures and may represent the major ancestral source contributed to them. A close genetic relationship was found between the Paleo-Inuit individual from the Saqqaq site in Greenland and the Syalakh-Belkachi population. 3-way models for Saqqaq individual results in ~64% – ~79% Syalakh-Belkachi-like admixture with the remaining from Kolyma_M_10.1kya and an East Asian source rich in Inland Northeast Asian ancestry (Zeng et al. 2023, sup. mat. p. 204).
In turn, the Saqqaq related ancestry persisted on either side of the Bering Straits among the ancient populations related to Eskimo-Aleut speakers. It is present at high levels in individuals from Old Bering Sea sites (~ up to ~43%) and in an individual from the classic Thule culture (up to ~59%), who is very similar to present-day Inuit. On the other hand a Native American-related source is present in passing models for all of these groups, which in the case of the Old Bering Sea culture suggests back-migration from the New World to Chukotka (Zeng et al. 2023, sup. mat. p. 212).
Saqqaq.SG ancestry persists along the Northern shores of the Sea of Okhotsk in a population from the Tokarev culture from ~3000 BP (Magadan_BA). This population is very similar to present-day Chukotko-Kamchatkans, and has very complex ancestry, with contributions from at least a Kolyma_M_10.1kya-related source and a Native American-related source on top of Saqqaq.SG ancestry; the remainder is drawn from East Asian-related sources (Zeng et al. 2023, sup. mat. p. 212).
Ancestry related to that found in Syalakh-Belkachi also persists in an Iron Age Yakutian (~2600BP) from the Middle Lena River Valley, who can be modeled as cladal with Yakutia_LNBA. This individual is extremely similar to present-day Yukaghirs and Nganasans (Zeng et al. 2023, sup. mat. p. 212)
While all ancient populations from Beringia and Arctic North America require a Saqqaq.SG-related source for passing models,suprisingly Ancient Athabaskans do not share in this pattern. The models indicate that ancient Athabaskans can be modeled without any ancestry from Saqqaq.SG (i.e., with Saqqaq.SG retained in the reference populations); rather, in a 3-way model where most ancestry comes from a Native-American-related and a Kolyma_M_10.1kya-related source, model p-values are maximized by a minor (~9-13%) contribution from a third source that may be either Cisbaikal_LNBA or the closely-related Ust_Kyakhta_14kya (Zeng et al. 2023, sup. mat. p. 213).
Yakutia_LNBA
Yakutia_LNBA, comprising individuals dating to ~4500-3200 years BP and from a region stretching from the Kan river valley in the Southeastern Krasnoyarsk region, to the Middle to Lower Lena valley in Central Yakutia and the Kolyma River basin in far Northeastern Yakutia close to Beringia, can be modeled as a three-way admixture between Syalakh-Belkachi (~50%), Transbaikal_EMN (~42%), and a source related to Amur River-related populations (~8%) (Zeng et al. 2023, sup. mat. p. 182).
The Yakutia_LNBA genetic cluster has strong correlation with the Uralic speaking populations. The recent comprehensive analysis shows that this ancestry distinguishes Uralic speaking populations from their non-Uralic speaking neighbours. The Uralic peoples derive the majority, and in some cases all of their East Asian-related ancestry, from this source (Zeng et al. 2023, sup. mat. p. 214-215)
Cisbaikal_LNBA
A model of Cisbaikal_LNBA as deriving most of its ancestry (~86%) from Ust_Kyakhta_14kya, with some ancestry from Inland Northeast Asians (~14%), is the only model passing with a high p-value. The origin of the Cisbaikal_LNBA is not clear and thought to be descended from an unsampled population that is related to but younger than Ust_Kyakhta_14kya (Zeng et al. 2023, sup. mat. p. 184).
Cisbaikal_LNBA ancestry, represented mostly by the individuals associated with Glazkov Culture, shows strong correlation with Yenisean speakers and constantly required as a source when modelling populations in the Yenisei Basin such as Yenisean speaking Kets, Uralic speaking Enets and Selkup and Turkic speaking populations (Tuvinian, Tofalar, Tubalar, Altaian, Altaian_Chelkan, Kakhass, Kakhass_Kachin, Shor, Shor_Mountain, and Todzin) which is not the case for other Turkic or Uralic peoples (Zeng et al. 2023, sup. mat. p. 219-220)
China_NEastAsia_Inland_EN
Represented by the Yumin (~8000 BP) individual. This ancestry seems to have significantly contributed to many later populations in Northeast Asia.
Jomon Ancestry
Represented by 8,000–3,000-year-old hunter-gatherer individuals in Japan. The oldest individual sampled to date is Higashimyo, from Kyushu, Japan. Populations associated with this ancestry contributed partially to present-day Japanese populations (Yang et al. 2022).
Southeast Asia
China_YellowRiver_N
Represented by the Neolithic individuals from the Yellow River region, China. This ancestry is associated with the Proto Tibeto-Sinitic languages and the major ancestral source that contributed to present day South and East Asian populations (Han Chinese, Tibetan, Korean, Japanese and etc.). The Neolithic individuals grouped under China_NEastAsia_Coastal_EN label are also closely related to the Neolithic Yellow River ancestry (Yang et al. 2022).
Fujian Ancestry
The ancestry described in Yang et. al, 2020 and 2021 , consisted of 4 groups based on the Neolithic era samples obtained from Fujian, China;
China_SEastAsia_Coastal_EN,
China_SEastAsia_Coastal_LN,
China_SEastAsia_Island_EN,
China_SEastAsia_Island_LN.
This ancestry is found in high levels in Austronesian populations and thought to be associated with Austronesian expansions. It is also one of the most significant ancestries shared in high levels among the present day East and Southeast Asians.
South Asia
Ancient Ancestral South Indian (AASI)
AASI is another “ghost” population, detected by indirect methods. Ancestry associated with the AASI lineage was found at low levels in almost all present-day Indian populations, particularly southern Indians. This ancestry has deep relationships with the other ancient East Eurasian ancestries and Onge or Irula populations frequenlty used as a proxy to represent this ancestry (Yang et al. 2022).
- Zeng, T. C., et al. (2023). Postglacial genomes from foragers across Northern Eurasia reveal prehistoric mobility associated with the spread of the Uralic and Yeniseian languages (pre-Print). BioRixv. https://doi.org/10.1101/2023.10.01.560332
- Yang, M., (2022). A genetic history of migration, diversification, and admixture in Asia. Hum Popul Genet Genom. 2(1):0001. https://doi.org/10.47248/hpgg2202010001
- Yang, M., et al. (2020). Ancient DNA indicates human population shifts and admixture in northern and southern China. Science 369,282-288. DOI:10.1126/science.aba0909
Further Reading
Near East
- The genetic structure of the world’s first farmers (2016)
- Genomic insights into the origin of farming in the ancient Near East (2016)
- The Demographic Development of the First Farmers in Anatolia (2016)
- Late Pleistocene human genome suggests a local origin for the first farmers of central Anatolia (2019)
- Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions (2019)
- The Genomic History of the Bronze Age Southern Levant (2020)
- Genomic History of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus (2020)
- The genetic history of the Southern Arc: A bridge between West Asia and Europe (2022)
- Spatial and temporal heterogeneity in human mobility patterns in Holocene Southwest Asia and the East Mediterranean (2023)
Europe
- Massive migration from the steppe was a source for Indo-European languages in Europe (2015)
- The genetic history of Ice Age Europe (2016)
- The Genetic History of Northern Europe (2017)
- The Genomic History of Southeastern Europe (2018)
- Understanding 6th-century barbarian social organization and migration through paleogenomics (2018)
- Corded Ware cultural complexity uncovered using genomic and isotopic analysis from south-eastern Poland (2020)
- Genetic ancestry changes in Stone to Bronze Age transition in the East European plain (2021)
- The Anglo-Saxon migration and the formation of the early English gene pool (2022)
- Population genomics of the Viking world (2022)
- Fine-scale sampling uncovers the complexity of migrations in 5th–6th century Pannonia (2023)
- Patrilocality and hunter-gatherer-related ancestry of populations in East-Central Europe during the Middle Bronze Age (2023)
- Genetic admixture and language shift in the medieval Volga-Oka interfluve (2023)
- A genetic history of the Balkans from Roman frontier to Slavic migrations (2023)
- Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers (2023)
- Population Genomics of Stone Age Eurasia (2024)
Africa
Eurasia
- Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe (2017)
- 137 ancient human genomes from across the Eurasian steppes (2018)
- Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads (2018)
- The first horse herders and the impact of early Bronze Age steppe expansions into Asia (2018)
- Reconstructing the Deep Population History of Central and South America (2018)
- The formation of human populations in South and Central Asia (2019)
- Y-chromosome haplogroups from Hun, Avar and conquering Hungarian period nomadic people of the Carpathian Basin (2019)
- Shifts in the Genetic Landscape of the Western Eurasian Steppe Associated with the Beginning and End of the Scythian Dominance (2019)
- A Dynamic 6,000-Year Genetic History of Eurasia’s Eastern Steppe (2020)
- Genetic insights into the social organisation of the Avar period elite in the 7th century AD Carpathian Basin (2020)
- The genomic origins of the Bronze Age Tarim Basin mummies (2021)
- Ancient genomic time transect from the Central Asian Steppe unravels the history of the Scythians (2021)
- The genetic origin of Huns, Avars, and conquering Hungarians (2022)
- Ancient genomes reveal origin and rapid trans-Eurasian migration of 7th century Avar elites (2022)
- Bronze and Iron Age population movements underlie Xinjiang population history (2022)
- Genetic population structure of the Xiongnu Empire at imperial and local scales (2023)
- Postglacial genomes from foragers across Northern Eurasia reveal prehistoric mobility associated with the spread of the Uralic and Yeniseian languages (2023)
East & Southeast Asia
- Ancient DNA indicates human population shifts and admixture in northern and southern China (2020)
- Genomic Insights into the Demographic History of Southern Chinese (2020)
- Ancient genomes from northern China suggest links between subsistence changes and human migration (2020)
- Ancient genomics reveals tripartite origins of Japanese populations (2021)
- Genomic insights into the formation of human populations in East Asia (2021)
- The deep population history of northern East Asia from the Late Pleistocene to the Holocene (2021)
- Human population history at the crossroads of East and Southeast Asia since 11,000 years ago (2021)
- A genetic history of migration, diversification, and admixture in Asia (2022)
- Human genetic history on the Tibetan Plateau in the past 5100 years (2023)