Ancient Populations
The following are categories of Paleolithic populations, their descriptions, and citations:
Dzudzuana
The Dzudzuana population represents a distinct lineage of Upper Paleolithic foragers in West Eurasia, identified through ancient DNA analysis from Dzudzuana Cave in Georgia (~26,000 years ago). Their genetic composition reveals a mixture of a deeply rooted Eurasian lineage and an enigmatic basal population, predating and potentially ancestral to early Neolithic farmers. Genomic analyses indicate that Dzudzuana-related ancestry significantly contributed to the genetic makeup of Neolithic populations in Anatolia and Europe, challenging earlier models that posited a strictly Near Eastern origin for early farmers. Comparative studies suggest a close genetic relationship between Dzudzuana and pre-agricultural groups in the Caucasus and the Zagros region, highlighting long-term interactions among foragers across these areas.
1.Lazaridis et al. (2018): “Paleolithic DNA from Caucasus reveals core of West Eurasian ancestry” – This paper explores the genetic legacy of the Dzudzuana population and its role in shaping early Neolithic groups.
2.Feldman et al. (2019): “Late Pleistocene human genome suggests a local origin for the first farmers of central Anatolia” – Discusses how Dzudzuana-related ancestry connects to later farming populations.
Ancient North Eurasians (ANE)
Ancient North Eurasians (ANE) were a genetically distinct population that thrived in Upper Paleolithic Siberia, with their ancestry primarily associated with the Mal’ta-Buret culture (~24,000 years ago). Genomic studies have demonstrated that ANE individuals were more closely related to Western Eurasians than to contemporary East Asians. Their genetic legacy is evident in later populations, including Indo-European steppe pastoralists (e.g., the Yamnaya culture) and Indigenous groups in the Americas, through their contribution to the Ancient Beringians. ANE ancestry remains prevalent in modern Central Asian, South Asian, and some Eastern European populations. Key individuals representing this ancestry include Mal’ta Boy (MA-1) and individuals from the Afontova Gora site.
1.Raghavan et al. (2014): “Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans” – One of the key papers identifying the Ancient North Eurasian population through the genome of Mal’ta Boy (MA-1).
2.Lazaridis et al. (2016): “Genomic insights into the origin of farming in the ancient Near East” – Provides further evidence of ANE contributions to later Eurasian populations.
Ancient Northern East Asians (ANEA)
Ancient Northern East Asians (ANEA) were a distinct genetic group that emerged in Pleistocene Siberia and Northeast Asia, diverging from other early East Asian-related populations. Genetic evidence places ANEA ancestry in Neolithic hunter-gatherers from sites such as Devil’s Gate Cave (~7,700 years ago) and in later populations speaking Tungusic, Mongolic, and Turkic languages. ANEA ancestry also played a crucial role in the genetic composition of Native American populations, particularly through ancestral groups in the Amur and Beringian regions. This lineage remains strongly represented in modern Northeast Asians, including the Han Chinese, Mongols, Koreans, and Siberian ethnic groups such as the Evenks and Nivkh.
1.Qiaomei Fu et al. (2016): “The genetic history of Ice Age Europe” – Covers the genetic differentiation of early Eurasian populations, including those in northern East Asia.
2.Gakuhari et al. (2020): “Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations” – Examines the deep ancestry of northern East Asian populations in Siberia and Japan.
Ancient Southern East Asians (ASEA)
Ancient Southern East Asians (ASEA) represent a basal East Asian lineage that contributed to the genetic makeup of present-day Southeast Asian and Austronesian-speaking populations. Genetic studies suggest that ASEA diverged early from other East Asian-related groups and share deep ancestral ties with pre-Neolithic populations in southern China, mainland Southeast Asia, and Taiwan. Modern proxies for ASEA ancestry include Austroasiatic-speaking populations such as the Mlabri and Maniq, as well as Indigenous groups from the Philippines, including the Aeta and Agta. The genetic imprint of ASEA ancestry is also evident in Negrito populations across Southeast Asia, underscoring the complex demographic history of the region.
1.McColl et al. (2018): “Ancient Genomics reveals four prehistoric migration waves into Southeast Asia” – One of the most comprehensive studies on ASEA and their contributions to modern Southeast Asian populations.
2.Lipson et al. (2018): “Ancient genomes document multiple waves of migration in Southeast Asian prehistory” – Highlights the genetic structure and migrations of ASEA populations.
Ancient Rainforest Hunter-Gatherers (Ancient RHG)
The Ancient Rainforest Hunter-Gatherers (RHG) represent an early-diverging lineage of humans that adapted to the dense tropical rainforests of Central Africa. Genomic analyses indicate that their closest modern genetic relatives are Mbuti and Biaka Pygmies, with a deep divergence from other African populations estimated at over 50,000 years ago. Their genetic isolation has contributed to unique adaptations related to stature, metabolism, and immune function, allowing them to thrive in rainforest environments. Due to the scarcity of ancient DNA from Central Africa, much of what is known about RHG ancestry comes from modern genomic studies, which reveal limited but detectable gene flow from Bantu agriculturalists in the past few thousand years. Their adaptation strategies included specialized dietary practices, metabolic adjustments, and cultural innovations suited to the challenges of rainforest foraging.
Citations:
1.Patin et al. (2014): “The impact of agricultural emergence on the genetic history of African rainforest hunter-gatherers and agriculturalists”
2.Hsieh et al. (2016): “Model-based analyses of whole-genome data reveal a complex evolutionary history involving archaic introgression in African hunter-gatherers”
South African Hunter-Gatherers
South African Hunter-Gatherers represent one of the most ancient surviving human lineages, ancestral to modern Khoe-San populations. Their genetic divergence from other African groups is estimated to have occurred approximately 200,000–300,000 years ago, making them one of the oldest distinct branches of modern humans. Archaeological evidence, including the Blombos Cave artifacts (~75,000 years ago), demonstrates early symbolic behavior, complex tool-making, and social structures indicative of advanced cognitive abilities. These populations traditionally maintained a foraging lifestyle in the Kalahari and surrounding regions, with genetic adaptations to arid environments and a diet rich in tubers and wild game. The modern San populations retain high levels of genetic diversity, reflecting their deep evolutionary history and long-standing genetic continuity.
1.Schlebusch et al. (2017): “Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago”
2.Pickrell et al. (2012): “The genetic prehistory of southern Africa”
Jomon
The Jomon people were a pre-agricultural population that inhabited prehistoric Japan (~14,000–300 BCE), known for their distinctive pottery traditions, hunter-gatherer subsistence strategies, and genetic distinctiveness from later agricultural societies. Ancient DNA analysis suggests that the Jomon carried a genetic profile intermediate between Ancient East Asians and more deeply diverged Southeast Asian populations, potentially reflecting early coastal migration routes from mainland Asia. Their genetic legacy remains most prominent in modern Ainu populations of Hokkaido, who retain a higher proportion of Jomon ancestry compared to other Japanese groups. However, with the arrival of Yayoi agriculturalists (~1000 BCE), the genetic landscape of Japan was significantly reshaped, leading to admixture between Jomon and incoming East Asian farming populations.
1.Gakuhari et al. (2019): “Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations”
2.Jinam et al. (2021): “Discerning the origins of the Japanese people from genome-wide data”
Basal East Asians (Onge)
The Onge people of the Andaman Islands are considered one of the closest modern representatives of an early-diverging branch of Basal East Asian ancestry. Genetic studies indicate that the Onge split from other East Asian-related populations before the differentiation of northern and southern East Asian lineages, possibly over 40,000 years ago. Their genetic distinctiveness is marked by low Neanderthal introgression and high genetic drift, likely due to long-term island isolation[1]. Recent research has revealed that the Zagros Neolithic Farmers (~10,000 BP, Iran) carried approximately 10% Basal East Asian ancestry, best represented by Onge-related genetic components. Similarly, the Caucasus Hunter Gatherers were found in the same study to have around 8% Basal East Asian ancestry as represented by Tianyuan. This suggests that early interactions between populations from South Asia and the Near East may have played a more significant role in shaping early Eurasian genetic diversity than previously assumed[2].
The Onge, along with other Andamanese populations, exhibit genetic continuity with early Hoabinhian hunter-gatherers of Southeast Asia, making them key to understanding early human migrations into tropical Asia and the Pacific.
1.Reich et al. (2009): “Reconstructing Indian Population History”
2.Lazaridis et al. (2018): “Paleolithic DNA from the Zagros reveals the genetic history of early farmers”
The following are reconstructed Paleolithic populations:
Ancient East African
The Ancient East African population represents an early-diverging lineage of hunter-gatherers and pastoralists who inhabited East Africa before and during the emergence of food production. Ancient DNA from Mota (~4,500 BP, Ethiopia) and Pastoral Neolithic populations (~3,000 BP) reveals a genetic continuum between indigenous foragers and Afroasiatic-speaking pastoralists. These populations played a crucial role in shaping the genetic landscape of modern Nilotic and Cushitic-speaking groups.
Unlike many other sub-Saharan Africans, Ancient East Africans exhibit Eurasian gene flow dating back ~7,000 years, likely introduced through early interactions with Near Eastern groups. However, Mota’s genome lacks this West Eurasian component, suggesting that admixture with Middle Eastern populations occurred after his time. Pastoral Neolithic individuals show ancestry from both indigenous East African foragers and incoming Afroasiatic-speaking herders, marking the transition to food production in the region.
1.Lucas-Sánchez et al. (2021): “Population history of North Africa based on modern and ancient genomes”
2.Shriner et al. (2018): “Genetic ancestry of Hadza and Sandawe peoples reveals ancient population structure in Africa”
Ancient North African
The Ancient North African population is inferred by analyzing the genetic structure of early North African groups before significant Eurasian admixture. The Iberomaurusian culture (~15,000 BP, Morocco, Algeria, and Tunisia) provides the earliest ancient DNA evidence from the region, particularly the Taforalt individuals (~15,000 BP, Morocco). Genetic studies show that Iberomaurusians carried a distinct indigenous North African component (~60-70%), mixed with some Dzudzuana-related ancestry (~30-40%). After removing Eurasian influences, the remaining ancestry represents a deep-rooted North African lineage likely dating back to the earliest modern human dispersals into the region.
This lineage is genetically distinct from both sub-Saharan African and West Eurasian populations, positioning it as an intermediate population between early Africans and Eurasians. Some genomic studies suggest a distant connection to Basal Eurasians, possibly due to an ancient back-migration from the Levant (~40,000 BP), followed by a long period of genetic isolation. Ancient North Africans show genetic continuity with later Berber-speaking populations, though modern North Africans carry additional Eurasian gene flow from Neolithic and historical periods.
1.Lucas-Sánchez et al. (2021): “Population history of North Africa based on modern and ancient genomes”
2.Henn et al. (2012): “Genomic ancestry of North Africans supports back-to-Africa migrations”
Ancient West African
The Ancient West African population, including the Shum Laka individuals (~8,000 BP, Cameroon), represents a highly diverse and ancient lineage of hunter-gatherers and early agriculturalists. Genetic analysis of these individuals reveals a Basal West African lineage that diverged from other African populations over 250,000 years ago. This lineage predates the emergence of Bantu-speaking populations and does not directly contribute to modern Niger-Congo-speaking groups.
The Shum Laka genome suggests an ancestral structure consisting of ~70% deep-rooted West African lineage and ~30% archaic introgression from an unknown hominin, indicating interbreeding with archaic humans within Africa. Unlike East and North African populations, Ancient West Africans show little evidence of Eurasian gene flow, reinforcing that their evolution occurred largely independently until later agricultural expansions.
This Basal West African lineage provides crucial insights into African genetic diversity. While not direct ancestors of modern Bantu populations, these early West Africans influenced the genetic makeup of modern Niger-Congo-speaking groups, particularly Yoruba and Mende, who also exhibit admixture from later migrations (~3,000 BP). Ancient West African ancestry is also a major genetic contributor to the African Diaspora, particularly among populations in the Americas with West African heritage.
1.Lipson et al. (2020): “Ancient West African foragers in the context of African population history” – Shum Laka Study
2.Schlebusch & Jakobsson (2018): “African genomic diversity and its implications for human evolution”
The following are Neolithic populations:
Amur River Hunter-Gatherers
The Amur River Hunter-Gatherers were a foraging population that occupied the Russian Far East and northeastern China during the Late Pleistocene and early Holocene. Genomic analysis suggests that they were closely related to Ancient Northern East Asians (ANEA) and significantly contributed to Siberian, Tungusic, and Native American populations[1].
Archaeological remains from Devil’s Gate Cave (~7,700 BP) in the Russian Far East indicate that these hunter-gatherers exhibited genetic continuity with later Jomon, Koreanic, and Tungusic-speaking groups[2]. Their genetic profile is distinct from the Yellow River Neolithic farmers, reflecting an early divergence within East Asian populations[3].
1.Sikora et al. (2019): “The population history of northeastern Siberia since the Pleistocene”
2.He et al. (2023): “Extensive ethnolinguistic diversity at the crossroads of North China and South Siberia reflects multiple sources of genetic diversity”
3.Bennett et al. (2024): “Reconstructing the Human Population History of East Asia through Ancient Genomics”
Anatolia Neolithic Farmers
The Anatolia Neolithic Farmers (~8,500–7,000 BP) were among the earliest groups to practice agriculture, playing a central role in the Neolithic Revolution. Their genetic influence extended across Europe and the Mediterranean, forming the primary ancestral component of early European farmers[1].
Genetic studies show that Anatolia Neolithic Farmers descended from a Dzudzuana-like forager population, with additional Levantine and Iranian admixture[2]. Their farming economy expanded via the Aegean route, replacing most hunter-gatherer populations in Europe while mixing with Western Hunter-Gatherers (WHG). They later contributed to Caucasus populations and Indo-European steppe pastoralists[3].
Citations:
1.Liu et al. (2021): “Insights into human history from the first decade of ancient human genomics”
2.Schurr & Pipes (2011): “The prehistory of Mongolian populations as revealed by studies of osteological, dental, and genetic variation”
3.Nägele et al. (2022): “Ancient genomic research-From broad strokes to nuanced reconstructions of the past”
Ancient Australians
The Ancient Australians were the first modern humans to arrive in Australia (~65,000 BP), carrying a deeply diverged lineage linked to Papuan and Melanesian populations. Genetic analyses place them within the Australasian branch of modern humans, exhibiting significant Denisovan admixture (~4–6%)[1].
Early skeletal remains such as Mungo Man (~40,000 BP) indicate long-term genetic continuity, reflecting adaptation to arid environments and isolation from other human populations for tens of thousands of years. Modern Aboriginal Australians remain one of the world’s oldest continuous cultures, retaining ancestral genetic signals dating back to their initial migration[2].
1.Bennett et al. (2024): “Reconstructing the Human Population History of East Asia through Ancient Genomics”
2.Rogers (2016): “Understanding ancient human population genetics of the eastern Eurasian steppe through mitochondrial DNA analysis”
Baikal Hunter-Gatherers
The Baikal Hunter-Gatherers (~9,000–6,000 BP) were a distinct foraging population in the Lake Baikal region of Siberia, closely related to Ancient Northern East Asians (ANEA). Genetic studies show that they contributed to later Siberian groups, such as Yukaghirs and Evenks, as well as to Native American populations[1].
Their genetic structure was separate from Beringian and Yakutian hunter-gatherers, indicating a complex population history in ancient Siberia. The region witnessed significant demographic shifts, with influences from both West Eurasian and East Asian populations over time[2].
1.de Barros Damgaard et al. (2018): “The first horse herders and the impact of early Bronze Age steppe expansions into Asia”
2.Sikora et al. (2019): “The population history of northeastern Siberia since the Pleistocene”
Caucasus Hunter-Gatherers (CHG)
The Caucasus Hunter-Gatherers (~13,000–10,000 BP) were an isolated Upper Paleolithic population inhabiting the Caucasus Mountains. Their genetic legacy contributed significantly to later Indo-European, Iranian, and South Asian gene pools[1].
CHG individuals were genetically distinct from Anatolia Neolithic Farmers and Eastern European Hunter-Gatherers (EHG) but later merged with them to form the Yamnaya steppe pastoralists (~5,000 BP), a key group in the spread of Indo-European languages[2]. Their DNA also contains a Basal Eurasian component, indicating early connections to Near Eastern populations[3].
1.Nägele et al. (2022): “Ancient genomic research-From broad strokes to nuanced reconstructions of the past”
2.Liu et al. (2021): “Insights into human history from the first decade of ancient human genomics”
3.He et al. (2021): “Genomic insights into the differentiated population admixture structure and demographic history of North East Asians”
East African Hunter-Gatherers (Mota/Bayira)
The Mota individual (~4,500 BP, Ethiopia) represents one of the earliest sequenced genomes from an East African forager. His genetic profile confirms that West Eurasian admixture in East Africa occurred later (~3,000 BP), supporting the hypothesis that the Afroasiatic language family’s genetic input from the Near East postdates his existence[1].
Genetic comparisons reveal that Mota shares ancestry with modern Omotic, Cushitic, and Nilotic-speaking populations, but he predates the expansion of pastoralism in East Africa[2]. His genome also suggests that pre-Bantu East African foragers had deep, indigenous ancestry with minimal external influence at that time.
1.Gallego Llorente et al. (2015): “Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent”
2.Skoglund et al. (2017): “Reconstructing Prehistoric African Population Structure”
East African Pastoral Neolithic
The East African Pastoral Neolithic (~3,000 BP) represents the transition from hunter-gatherer societies to pastoralist economies in Kenya and Tanzania. This period saw the introduction of cattle, sheep, and goats, likely introduced by Afroasiatic-speaking herders from the Near East[1].
Genetic studies show that Pastoral Neolithic populations descended from both indigenous East African foragers and Near Eastern herders, marking the first major period of admixture in the region. Their genetic legacy is especially pronounced in modern Nilotic and Cushitic pastoralist groups, who retain significant ancestry from these populations[2].
1.Prendergast et al. (2019): “Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa”
2.Wang et al. (2022): “Genomic insights into the formation of human populations in East Africa”
Eastern European Hunter-Gatherers (EHG)
The Eastern European Hunter-Gatherers (EHG) (~8,000 BP) occupied the regions of modern-day Russia and Ukraine, forming a significant component of Proto-Indo-European steppe ancestry[1].
Genetic studies reveal that EHG individuals were a mixture of Ancient North Eurasians (ANE) and Western Hunter-Gatherers (WHG), distinguishing them from purely Western or Siberian populations. Their genetic legacy is particularly evident in the Yamnaya steppe pastoralists (~5,000 BP), who later played a key role in the spread of Indo-European languages into Europe and South Asia[2].
1.Mathieson et al. (2018): “The Genomic History of Southeastern Europe”
2.Haak et al. (2015): “Massive migration from the steppe was a source for Indo-European languages in Europe”
Hoabinhian Hunter-Gatherers
The Hoabinhian people (~10,000–4,000 BP) were an early foraging population in Southeast Asia, preceding the arrival of Neolithic rice farmers from China.
Genomic studies indicate that modern Southeast Asians, particularly Austroasiatic-speaking and Negrito groups, retain significant Hoabinhian ancestry. While they were gradually replaced by Neolithic farmers, genetic evidence suggests they intermixed rather than being completely displaced[1].
1.McColl et al. (2018): “Ancient Genomics reveals four prehistoric migration waves into Southeast Asia”
Iberomaurusians
The Iberomaurusians (~15,000 BP, North Africa) were one of the earliest known populations in the Maghreb, preceding the spread of agriculture into the region.
Ancient DNA from Taforalt (~15,000 BP, Morocco) reveals that Iberomaurusians carried a distinct North African genetic component (~70%), mixed with Dzudzuana-related ancestry (~30%). Their legacy persists in modern Berber-speaking populations, although later gene flow from the Near East and Europe during the Neolithic and Bronze Age reshaped the genetic landscape of North Africa[1].
1.van de Loosdrecht et al. (2018): “Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations”
Jomon
The Jomon lineage is estimated to have diverged from other East Asian populations around 22,000–25,000 years ago. Genetic analyses of Jomon remains, such as the IK002 individual, confirm that the Jomon formed a distinct clade separate from later East Asians[1].
A 2021 study by Adachi et al. estimated that the Jomon genetic divergence occurred approximately 22,000 years ago, predating major migrations that shaped the genetic structure of mainland East Asians[2]. Other studies suggest that the Jomon carried a unique genetic signature, possibly influenced by Northern Eurasian and early East Asian populations, but remained highly isolated from continental populations for thousands of years[3].
The Jomon divergence is further supported by their high levels of genetic drift, which resulted from their long-term isolation on the Japanese archipelago. This distinct lineage remained largely intact until the arrival of Yayoi agriculturalists (~3,000 BP), which significantly altered the genetic landscape of Japan[4].
1.Gakuhari et al. (2020): “Ancient Jomon genome sequence analysis sheds light on migration patterns of early East Asian populations”
2.Adachi et al. (2021): “Ancient genomes from the initial Jomon period: new insights into the genetic history of the Japanese archipelago”
3.Kanzawa-Kiriyama et al. (2019): “Late Jomon male and female genome sequences from the Funadomari site in Hokkaido, Japan”
4.Gakuhari et al. (2019): “Jomon genome sheds light on East Asian population history”
Ancient Melanesians
The Ancient Melanesians (~50,000 BP) were among the earliest modern human populations to settle in the Pacific, forming a deeply diverged lineage within Australasian populations. They descend from the first Out of Africa migrants who traveled into Southeast Asia and Oceania, sharing ancestry with Ancient Australians and Papuans.
One of their most defining genetic characteristics is their high Denisovan admixture (~4–6%), which contributed to adaptations to tropical environments[1]. Ancient Melanesians also played a significant role in shaping the genetic makeup of Polynesians, Micronesians, and Indigenous groups of New Guinea, maintaining a distinct evolutionary trajectory due to their geographic isolation.
1.Reich et al. (2011): “Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania”
Natufian Hunter-Gatherers
The Natufians (~12,500–9,500 BP, Levant) were one of the first semi-sedentary foraging societies, marking the transition from hunter-gathering to early agriculture. They played a crucial role in the Neolithic farming revolution in the Fertile Crescent, cultivating early forms of wheat and barley.
Genetically, Natufians show ancestral links to North Africans and Levantine populations, sharing some Iberomaurusian-like ancestry (~25–30%). Unlike later Anatolian Neolithic Farmers, they lacked significant Basal Eurasian ancestry, suggesting a distinct genetic history[1]. Their cultural innovations led to the rise of Pre-Pottery Neolithic societies (~9,000 BP), which expanded into Europe and North Africa.
1.Lazaridis et al. (2016): “Genomic Insights into the Origin of Farming in the Ancient Near East”
North American Hunter-Gatherers
The North American Hunter-Gatherers (~14,000–5,000 BP) were the first populations to settle North America, descending from Ancient Beringians and Northeast Asian foragers. Early Paleoindian cultures, such as the Clovis tradition (~13,000 BP), are characterized by their distinctive stone tools and evidence of early megafauna hunting[1].
Genomic studies of Anzick-1 (~12,600 BP, Montana) reveal that Clovis individuals were closely related to South American hunter-gatherers, suggesting a rapid and early expansion of Paleoindian groups across the continent[2]. However, recent ancient DNA samples from Southwest Ontario (~4,500 BP) provide additional insights into North American genetic diversity, showing that these populations retained distinct ancestral lineages alongside Clovis-related groups[3]. The Ontario samples exhibit affinities to both earlier Paleoindian populations and later Indigenous groups, confirming a complex and regionally diverse genetic landscape in prehistoric North America.
By ~6,000 BP, additional migrations introduced new waves of Northeast Asian ancestry, further shaping the genetic diversity of Native American populations[4]. The combination of Clovis-related ancestry, early Beringian input, and later gene flow from Northeast Asia highlights the multi-layered demographic history of ancient North America.
1.Chatters et al. (2024): “Mammoth featured heavily in Western Clovis diet”
2.Raghavan et al. (2015): “Genomic evidence for the Pleistocene and recent population history of Native Americans”
3.Scheib et al. (2018): “Ancient human parallel lineages within North America contributed to a coastal expansion”
4.Grugni et al. (2019): “Analysis of the human Y-chromosome haplogroup Q characterizes ancient population movements in Eurasia and the Americas”
South American Hunter-Gatherers
The South American Hunter-Gatherers (~13,000 BP–present) were the first populations to inhabit South America, with genetic origins tracing to both Ancient North Eurasian (ANE) and Ancient Northern East Asian (ANEA) sources.
The Monte Verde site (~14,500 BP, Chile) provides evidence of early human settlement in South America, supporting a coastal migration route rather than an overland passage through North America. Ancient genomes from Lagoa Santa (~10,000 BP, Brazil) reveal a distinct genetic lineage, different from later Andean and Amazonian populations, suggesting an earlier, now-extinct migration wave[1].
1.Posth et al. (2018): “Reconstructing the Deep Population History of Central and South America”
Southeast Asian Neolithic Farmers
The Southeast Asian Neolithic Farmers (~4,000 BP) were early rice and millet cultivators whose migration across Mainland and Island Southeast Asia reshaped the region’s demographic landscape.
These farmers originated from Neolithic Yangtze River cultures (~6,000 BP) and expanded southward into Vietnam, Thailand, and the Philippines, significantly influencing Austroasiatic, Tai-Kadai, and Austronesian-speaking populations[1]. Their arrival triggered a demographic transition, leading to a mixing or replacement of Hoabinhian hunter-gatherers and forming the ancestral core of modern Southeast Asians.
1.McColl et al. (2018): “Ancient Genomics Reveals Four Prehistoric Migration Waves into Southeast Asia”
Western Hunter-Gatherers (WHG)
The Western Hunter-Gatherers (~8,000–5,000 BP, Europe) were the dominant Mesolithic foragers of Western and Central Europe, preceding the arrival of Neolithic farmers.
They were genetically distinct from Eastern European Hunter-Gatherers (EHG) but later admixed with them and with Anatolian Neolithic Farmers. Their ancestry is best observed in Mesolithic individuals such as Loschbour (~8,000 BP, Luxembourg) and La Braña (~7,000 BP, Spain)[1]. WHG populations contributed to later Bell Beaker and Neolithic European populations, though they were eventually partially replaced by incoming Indo-European pastoralists (~3,000 BP).
1.Haak et al. (2015): “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe”
Yakutia Hunter-Gatherers
The Yakutia Hunter-Gatherers (~6,000 BP, Siberia) were a cold-adapted population in northeastern Siberia, contributing to modern Indigenous Siberians and Native Americans.
They were genetically related to Ancient Northern East Asians (ANEA) and Baikal hunter-gatherers, later mixing with Turkic and Mongolic migrants to form the Yakut and Evenki peoples[1].
1.Sikora et al. (2019): “The Population History of Northeastern Siberia Since the Pleistocene”
Yellow River Neolithic Farmers
The Yellow River Neolithic Farmers (~7,000 BP, China) were early millet cultivators, forming the ancestral foundation of Han Chinese and Sino-Tibetan populations.
The Longshan culture (~5,000 BP) was a key late Neolithic civilization in northern China, known for its advanced pottery, early urbanization, animal domestication, and the development of complex social structures. Genetic and archaeological evidence suggests that Longshan farmers assimilated or displaced earlier Amur River and Hoabinhian hunter-gatherers, ultimately shaping the ancestral genetic profile of modern East Asians[1].
Citations:
1.Yang et al. (2020): “Ancient DNA Reveals the Genetic Impact of the Longshan Culture in China”
Zagros Neolithic Farmers
The Zagros Neolithic Farmers (~10,000 BP, Iran) were one of the earliest agricultural populations, distinct from Anatolian Neolithic Farmers but closely related to later Middle Eastern and South Asian groups.
Unlike Anatolian farmers, they carried higher levels of Basal Eurasian ancestry, making them genetically more distant from European populations. Over time, their genetic legacy contributed to the Indus Valley Civilization (~5,000 BP) and later Iranian and South Asian populations. By ~3,000 BP, they admixed with Steppe pastoralists, influencing Indo-Aryan migrations into South Asia[1].
1.Lazaridis et al. (2016): “Genomic Insights into the Origin of Farming in the Ancient Near East”
Reconstructed Neolithic samples:
Ancient Ancestral South Indian (AASI)
The Ancient Ancestral South Indians (AASI) represent the indigenous forager populations of South Asia before the arrival of Neolithic farmers and Indo-Aryan migrants. Modern South Indian tribal groups, such as Paniya and Irula, retain high proportions of AASI ancestry, providing a genetic model for reconstructing these early populations[1].
Genomic evidence suggests that AASI diverged from other non-African populations ~50,000–60,000 years ago, shortly after the initial migration Out of Africa. AASI groups were not homogeneous, but rather consisted of diverse foraging populations across India and Sri Lanka. By ~9,000 BP, Neolithic farmers from the Iranian Plateau (Zagros Neolithic Farmers) migrated into South Asia, mixing with AASI populations. Later, Indo-Aryan migrations (~3,500 BP) further reshaped the genetic landscape, reducing the relative proportion of AASI ancestry in North India.
The Harappan Civilization (~4,500–1,900 BCE) individuals show a significant AASI component, mixed with Zagros Neolithic ancestry, suggesting that AASI groups were partially integrated into early urban societies before later Indo-Aryan expansions[2]. Today, AASI ancestry is strongest in South Indian tribal and Austroasiatic-speaking groups, particularly in the Dravidian-speaking populations of the Indian subcontinent.
1.Narasimhan et al. (2019): “The Formation of Human Populations in South and Central Asia”
2.Shinde et al. (2019): “Ancient DNA from the Harappan Civilization Provides Insights into South Asian Ancestry”
Ancient Rainforest Hunter-Gatherers (Ancient RHG)
The Ancient Rainforest Hunter-Gatherers (Ancient RHG) represent an early forager population that inhabited Central Africa’s dense rainforest ecosystems, genetically reconstructed using modern Mbuti and Biaka Pygmies as well as ancient DNA from Shum Laka (~8,000 BP, Cameroon).
Rainforest adaptation among these populations dates back at least 20,000 years, supported by evidence of short stature, high disease resistance, and metabolic specializations linked to life in dense forests. Genetic studies suggest that Ancient RHG diverged from other sub-Saharan Africans ~60,000 years ago, forming a distinct lineage before the Bantu expansion (~3,000 BP) dramatically reshaped Africa’s genetic landscape[1].
Ancient DNA from Shum Laka indicates that modern Central African Pygmy groups (Mbuti and Biaka) descend from a population closely related to Ancient RHG but exhibit later Bantu and other external gene flow[2]. Today, remnants of Ancient RHG ancestry persist in Pygmy groups, though they have been largely displaced by Bantu-speaking agriculturalists.
1.Patin et al. (2014): “The Impact of Agriculture on the Genetic History of African Rainforest Hunter-Gatherers”
2.Lipson et al. (2020): “Ancient West African Foragers in the Context of African Population History”
Ancient West African
The Ancient West African population represents a genetically reconstructed hunter-gatherer and pre-Bantu group, modeled using ancient DNA from Shum Laka (~8,000 BP, Cameroon) and modern Yoruba and Mende populations.
Ancient West Africans were highly diverse, with distinct regional subgroups that preceded the spread of Bantu languages and agriculture (~3,000 BP). The Shum Laka individuals reveal a genetic profile that includes ~70% West African ancestry and ~30% admixture from an unidentified archaic hominin, suggesting possible interbreeding with an unknown hominin lineage within Africa[1].
Early West African groups likely contributed to the Niger-Congo-speaking populations that later expanded across Africa with the Bantu migration. Some West African populations also carry traces of Ancient North African or Saharan ancestry, reflecting early trans-Saharan interactions[2].
1.Lipson et al. (2020): “Ancient West African Foragers in the Context of African Population History”
2.Skoglund et al. (2017): “Reconstructing Prehistoric African Population Structure”
Ancient Nilote
The Ancient Nilote population is a genetically reconstructed profile of early Nilotic-speaking pastoralists, modeled using modern Dinka and other Nilotic-speaking populations of South Sudan.
Nilotic populations exhibit deep genetic divergence from other East African groups, dating back at least 20,000–30,000 years, likely originating in the central Nile Valley or Great Lakes region. Unlike surrounding foragers, Ancient Nilotes developed early pastoralist traditions, specializing in cattle herding and adopting a distinct subsistence economy[1].
The East African Pastoral Neolithic (~3,000 BP) represents a crucial transition, as early Nilotic groups expanded southward into Kenya, Uganda, and Tanzania, spreading herding practices. Despite maintaining strong genetic continuity from Ancient East Africans, Nilotic-speaking groups absorbed limited Afroasiatic (Cushitic) ancestry due to interactions with pastoralists from the Horn of Africa[2].
1.Prendergast et al. (2019): “Ancient DNA Reveals a Multistep Spread of the First Herders into Sub-Saharan Africa”
2.Wang et al. (2022): “Genomic Insights into the Formation of Human Populations in East Africa”