Q-L232 is a human Y-chromosome DNA haplogroup that is found primarily in Central Asia, Siberia and the Americas. It is also present in almost all Afghan and even many South Asian groups. Especially among some Yusufzai and Swat groups it can reach a high frequency. Afghans belong to a diverse set of Q clades, which often is of very different origins. Some of the oldest Q in Afghanistan could predate even Indo-Europeans, but much of it arrived likely later with Indo-Iranians in the Bronze Age and Turco-Mongols in the historic/medieval period, who received their Q lines from older Siberian and Inner Asian Mountain Corridor (IAMC) populations. These substrate populations rich in Y-DNA Q remain in many aspects very mysterious and only thanks to ancient DNA we became aware of their contribution to the gene pool of modern-day Afghans. Afghans not just have Y-DNA from these ancient Central Asians, but also some autosomal DNA and mtDNA. It is even possible that some terms of their extinct languages survived as loanwords in Indo-Iranian languages and Burushaski.
Especially important in the context of Q-L232 in Afghans is the Inner Asian Mountain Corridor (IAMC), which as an ancient exchange route ranging from the Altai Mountains in Siberia to the Hindu Kush played a central role in the ancient history and diffusion of Q-L232. It linked the mammoth steppes of North Eurasia with South Eurasia and connected in the Neolithic and Bronze Age various fertile valleys rich in animal, metal, stone and water resources. The mammoth steppes and valleys of Siberia would act as a refugium during the last Late Glacial Maximum (LGM) around 20.000-30.000 years ago when much of the Inner Asian Mountain corridor including the Pamir and Tian Shan Mountains were covered in glaciers. But later after massive deglaciation around 15.000 B.C we see much of the mountain valleys becoming accessible and via the Inner Asian Mountain corridor much of the arid regions of South Central Asia would later be repopulated after the Late Glacial Maximum. But the Inner Asian Mountain corridor also received genetic admixture from people arriving after the Ice Age in the region from the south, northwest, and east. In the southern part of the IAMC region groups from the Iranian Plateau rich in Neolithic Iranian admix arrived in the Neolithic. They would also bring new plants, animals, and technologies to the region. Most northeastern regions would receive East Asian admix already very high in Neolithic samples of the Altai region. From the northwest we see in the early Neolithic admix from EHG-admixed groups of the Central Steppe and finally Yamnaya-like admixture brought by Indo-Europeans of the Afanasievo and Andronovo culture (Andronovo also brought EEF admix from Europe to the region)
Haplogroup Q is around 30.000 years old and a distant relative of Haplogroup R, which like Q is a descendant of Haplogroup P. Haplogroup P likely originated in North Eurasia and was found in the ancient Upper Paleolithic Yana site (Raghavan et al 2014). Yana people were part of a population ancestral to mammoth hunters rich in Haplogroup R and Q that roamed across the mammoth steppes of Siberia, Central Asia and parts of Europe during the Paleolithic. Autosomally this Paleolithic population appears to have passed on its genes mostly to the populations of Europe, South-Central Asia and pre-columbian Americas. In archaeogenetics, the term Ancient North Eurasian (ANE) was given to an ancestral component that represented this ancient group of mammoth hunters. ANE people would dominate much of Siberia and the IAMC region up to the Neolithic and in some regions even Bronze Age.
Q-L232 from the Mesolithic to late Neolithic
The oldest ancient Q found yet in archaeogenetics is from the Afontova-Gora-2 site on the Yenisei River Bank near Krasnojarsk (Raghavan et al 2014). He was dated to 15.000 B.C and Q1a1-Y570. He had a typical ANE autosomal DNA profile with 1.9-2.7% of the genome of Neanderthal origin. Despite of the site being located in Central Siberia, which today is associated with East Asian ancestry, the sample lacked East Asian ancestry. So we see later an at least partial replacement of much of the ANE-rich people with Q and R by populations richer in East Asian autosomal dna and uniparental markers.
Artifacts of Paleolithic Afontova-Gora
Another female sample from the AfontovaGora site (AfontovaGora 3) carried the derived rs12821256 allele associated with, and likely causal for, blond hair color, making Afontova Gora 3 the earliest individual known to carry this derived allele. The allele was also found in three later members of the largely ANE-derived Eastern Hunter-Gatherers populations from Samara, Motala and Ukraine c. 8000 B.C, suggesting that it originated in the Ancient North Eurasian population before spreading to western Eurasia. But only much later in the Bronze Age and in Europe blonde hair would become common and more frequent because of selection and genetic founder effects.
Paleo-American Q1b1-L53
Carriers of Q1b1-L53 but mainly Q1b1a-L54 would around 15.000 B.C. cross the Bering Strait from North Asia into the Americas over the Beringa land bridge. This bridge existed from 45,000 to 12,000 BCE but ice-free corridors developed along the Pacific coast and valleys of North America only around 16,500 – c. 13,500 B.C. We even have very old ancient genomes from North and South America with Q1b1a-L54 like for example Anzick-1 ( 10.000 B.C, Clovis culture/USA), Kennewick (6800 B.C, USA) and I11974 (9935 B.C, Southern Cone/Chile). But it is interesting to mention, that one sample from Nevada dated to 200 A.D was Q1b1b-YP4010 (also under Q1b1-L53), unlike most other ancient and modern Native Americans.
These Q-rich people from Siberia were ancestral to Native Americans and Q1b1a is today the most frequent haplogroup among native people of North and South America. Native Americans and these ancient Paleo Americans are also one of the group of people highest in ANE ancestry (up to 55%,) but they also carry high amounts of East Easian ancestry unlike many other ancient ANE populations rich in Y-DNA Q and R.
Q1a1-YP570 in East Siberia
But most ancient Q remained in Siberia and North Eurasia. We have ancient DNA evidence for Q being present from the Baltic to the Beringian Sea in the Mesolithic and Neolithic. After AfontovoGora the oldest Q is from the Kolyma site in Yakutia (East Siberia) dated to 7800 B.C. The sample carried Q1a1-Y570 like Afontova-Gora but already was very rich in East Asian ancestry and carried around 60-70% of East Asian-related ancestry. So it also represents one of the earliest samples with East Asian ancestry in North Eurasia. Generally, we see that Q1a1-YP570 hunter-gathers of Siberia like the Paleolithic ancestors of Native Americans mixed early with people of East Asian ancestry in East Siberia. So it is not surprising that Q1a1-YP570 shows up in the Neolithic Yangshao culture of North China, Bronze Age Slab Grave culture of Mongolia (,..) and historic era samples from Mongolia and Beringia. It is also one of the Q clades, which would later be assimilated by Steppe Iranics and Turko-Mongolics, so possibly present in modern-day Afghans especially the ones of Turkic and Mongolic origin.
Q1b2a-Y6802 in early Neolithic Tajikistan
One of the oldest Q samples, which likely is also relevant for modern Afghans is from the Tutkaul site (sample Tutkaul 1) , which is located in Dashti-Mazar, 70 km southeastwards from Dushanbe where the Vakhsh enters the Pulisangin Gorge. Tutkaul 1 was dated to 6419-6026 B.C and is for now also one of the oldest ancient samples from South Central Asia we have. He was Q1b2a-Y6802, which is around 15.000 years old and was also found in the much later Bronze Age Afanasievo and Corded Ware culture. Today we also have Pashtuns, Tajiks and South Asians under Q-Y68202. But it is unclear if Tutkaul 1 is directly ancestral to Afghan Q1b2a the TMRCA is very old and Q1b2a seemingly was present from East Europe to Central Asia already in the Neolithic if not earlier.
The autosomal profile of the sample was closely related to Upper Palaeolithic individuals from south-central Siberia (Afontova Gora 3 (AG3) and Mal’ta 1), and roughly contemporaneous to West Siberian hunter-gatherers also rich in Y-DNA Q and ANE ancetry. Contrary to West Siberian hunter-gatherers, Mesolithic Kolyma, and Paleo Americans, Tutkaul 1 did not carry an extra-eastern Eurasian ancestry but had ancestry related to Neolithic Iranians. It is likely that the ancestors of Tutkaul 1 mixed with Mesolithic Zarzian or even early Neolithic Iranian groups rich in West Asian ancestry (Posth et al. 2023). Most likely ANE-rich people arrived in Tajikistan and South Central Asia only around 10.000 B.C with earlier populations in the region having connections to the Iranian Plateau and West Asia,
Q1b and Q1a in hunter-gathers of Europe
One of the oldest Q samples was found in Baltic Mesolithic Hunter Gathers in Latvia. These hunter-gathers lacked East Asian ancestry and were rich in East European hunter gather ancestry. East European Hunter Gathers had a high amount of ANE ancestry, which likely was the source of that Q. The sample I4550 was Q1b-Y2700 and dated to 6583-6420 B.C. He belonged to a basal Q1b line and probably has no modern descendants but the presence of Q1b in Baltic Hunter Gathers shows that Q had a wide distribution in North Eurasia in the Mesolithic and was associated with West Eurasian ancestry. Q1b1a1b-L804 was also found in hunter-gathers of the Volosovo culture in Central Russia dated to around 4000 B.C. This Q clade is the closest to Paleo-American Q-M3 clade in Eurasia. It is closer to Native American clades than to all other ancient Neolithic and Mesolithic Q clades found yet. But the TMCRA of the shared ancestor is estimated to be around 13.000 B.C and possibly a bit older. Today Q1b1a1b-L804 is found in low frequencies in Scandinavia and NW Europe. Also surprisingly Q1b was found in Neolithic Sweden among Funnel Beakers dated to 3500-3700 B.C, which likely assimilated it from local HG populations. Q1a, which generally had a more eastern distribution, also shew up among hunter-gathers of the Volosovo culture in the Volga-Kama region and in HG cultures of Mesolithic North Russia. But unlike Q1b1a1b this Q1a seems to be extinct today and even rarer than Q1b1a1b-L804.
Q1b1b and Q1b1a3 in South Siberia/Cis-Baikal
We also have many samples from the Neolithic Cis-Baikal region dated around 3000-4000 B.C with Q1b1b-YP4010 or Q1b1a3-L330. Like much earlier Kolyma in East Siberia these samples were very rich in East Asian ancestry (around 70% East Asian and 30% ANE-related). These Q lines would later show up in the Iron Age and historic eras of Steppe groups like Saka, Alans, and Turkic. But it is likely that they were originally associated with speakers of Paleo-Siberian and Yenissei languages like Ket, who today fall mostly under Q1b1a3-L330. It is likely that modern-day Hazara and Turkics of Afghanistan will show up these Saka and Turco-Mongolic-related Q clades.
Q in Bronze Age Eurasia
(Proto)-Indo-European Q1b2a1a-FT380500
We also have not few Q samples from the Eurasian Steppe in both early Indo-European and Pre-Indo-European contexts. One of these clades under Q1b2a1a even looks like a Proto-Indo-European line. Q1b2a1a-FT380500 was around 2600 B.C and found in the Indo-European Afanasievo culture of the Altai mountains on the Yenissei river but also in the Corded Ware culture in a site in Bohemia dated to 2700 B.C. Q1b2a1a-FT380500 has a TMRCA estimated to be dated around 3000 B.C., what strongly points to a common and recent otigin of the Afanasievo and Corded Ware Q1b2a1a. Interestingly upstream Q1b2a1a-Y147687 is today found among Tajiks and Sarikoli but these clades are not direct descendants of these Afanasievo and Corded Ware lines . Q1b2a1a-Y147687 has a TMRCA dated to around 13.000 B.C so it is likely that Q1b2a1a-Y147687 was present both in PIEs of Eastern Europe and Pre-IEs of Central Asia.
Central Eurasian Q1b2a1a-FT239746
Two very interesting Q-FT239746 samples from the Bronze Age were found in very different cultures and regions. I11526 from Bronze Age Kyrgyzstan at the Aigyrzhal site on the Naryn valley was Q-FT239746. He was dated to around 2100 B.C and a mix of people from the Iranian Plateau, IAMC, and Central Steppe region but he lacked Indo-Iranian Steppe MLBA admixture pointing to a slightly later arrival of Indo-Iranian in the region. Aigyrzhal people were agro-pastoralists who mainly practiced pastoralism but we have found evidence of domestication of wheat and barley at the site. it is likely that the first Indo-Iranian settlers in Central Asia would mix with such kind of people and spread ancestry related to them further south into Afghanistan and South Asia.
Another Q-FT239746 sample was found in the Indo-Iranian Sintashta culture. The sample I1017 was dated to around 1850 B.C but he was a extreme genetic outliner not of Steppe MLBA origin but rather of Central Steppe, West Siberian HG and IAMC origin. He clustered closest to some Central Steppe Pre-Andronovo samples like Kumsay_EBA.
Central-Eurasian Steppe Q1b2b1b-L392
Another possible Eurasian Steppe Q1b2 line but rather Pre-IE Steppe than PIE is Q1b2b1b-L932, which was found in the North Caucasian Steppe (Steppe Maykop) and Kazakh Steppe (Kumsay_EBA) during the early Bronze Age around 3000 B.C. Later we see Q-L392 in outliners of the Indo-Iranian Andronovo culture of Siberia and the Inner Asian Mountain Corridor (IAMC) region. For example, I4784 from Kyzyl Bulak 1 in Kazakhstan and I6717 from the Krasnoyarsk region in Russia had Q1b2b1b-L932 and were both dated around 1500 B.C belonging to an Andronovo-related archaeological context. These samples also shared Pre-Indo-Iranian Central Steppe ancestry like in Early Bronze Age Kumsay.
BMAC and Bronze Age IAMC Q
Q was also in the Bronze Age very diverse in South Central Asia and the IAM region. Here we find it both in Indo-Iranian and Pre-Indo-Iranian contexts. For example, a Q sample was found in the BMAC city of Gonur. BMAC was mainly of Neolithic Iranian origin but had admixture from IAMC populations especially the Q samples, which often were easily detectable as genetic outliners. The Gonur sample I6119 (2025 B.C) was Q1b1a-M346 so not directly related to much earlier Tutkaul 1 from Tajikistan. These South-Central Asian Q clades would later be later by Indo-Iranians and migrate southwards with R1a-Z93 of Proto-Indo-Iranians. Later we also see in an Andronovo-related context Q1a2a2-F4531 in Kokcha, Uzbekistan (I8506 dated to 1600 B.C.). The sample had local BMAC-like admix so possibly he received his Q1a2a2 from a local group but a Steppe origin is also possible. Much of the clades found in Andronovo- and BMAC-related contexts show up today in Afghanistan and South Asia, what strongly points to Indo-Iranians assimilating in Central Asia earlier pre-Indo-Iranian groups and bringing these new Q clades to South Eurasia.
Q1b2b1b-FT377000, Q1b1a3-L330 and Q1a2a1-YP832 in the Okunevo culture
The Okunevo culture was an archaeological culture that existed in the southern Siberian Khakassia region of Russia during the Bronze Age. It is believed to have existed between approximately 2,700 BCE and 1,800 BCE. It is known for its sophisticated and fascinating artworks.
The Okunevo people were cattle-breeding pastoralists, but their recent ancestors and genetic relatives were still hunter-gathers along the various tributaries in these regions. It is assumed that the ancestors of the Okunev migrated from north to south. From a genetic standpoint Okunev were West-Siberian hunter gatherers with an extra affinity to East Asian populations, which made up about 30% of their admixture. In addition, some Okunevo samples show Afanasievo ancestry, both in their autosomal makeup (10-20%) as well as their uniparental markers (R1b-Z2103). But most Okunevo people fall under Q1b1a3-L330 and Q1b2b1b-FT377000 with sample samples also carrying Q1a2a1-YP832. Q1b1a3-L330 already shew up earlier in Cis-Baikal samples but Q1b2b1b–FT377000 shew up for the first time in ancient dna and is an underresearched Q clade. It also has not many if any direct modern descendants but is closest to Q1b2b1a-L527, which today is specific for North Europeans. Still it is unlikely that these European Q1b2b1a is directly related to Okunevo people, because the TMRCA is estimated to have lived around 13.500 B.C. So North Euro Q-L527 could be an EHG line of people like Volosovo or Baltic Sea hunter-gathers, that survived in North Europe. But considering the quite young TMRCA of Q1b2b1a-L527 (1100 B.C) it could arrive in North Europe. Q1a2a1-YP832 shew up for the first time in archaeogenetics in Okunevo and would later show up in Hunnic samples from the Tian Shan region.
The Okunev culture was later replaced by the Indo-Iranian Andronovo culture, although the Tagar culture shows some Okunev-like genetic ancestry. It is possible they influenced the typical art forms of the Saka to a degree.
Q in Iron Age Eurasia and modern Afghans
Q2b-Y1150 in Iron Age Swat and modern Afghans
Ancient Bronze Age to Iron Age samples
- I7493; likely Iron Age; Sappali Tepe; Uzbekistan; BMAC_o2; Q1b2-L275>Y1150 (xYP4500, YP3943)
- I13228; 1000-800 BC; Loebanr, Swat Valley; Pakistan; SPGT; Q1b2-L275>Y1150>BY186390
- I5400; 927-831 BC; Loebanr, Swat Valley; Pakistan; SPGT; Q1b2-L275>Y1150>BY186390
- I7718; 400-200 BC; Saidu Sharif, Swat Valley; Pakistan; Swat_H; Q1b2-L275>Y1150* (xY1144)
- DA127,214-528 AD;Aktas-Issyk; Kazakhstan, Tian Shan Hun_O;Q1b2-L275>Y1150>Z5901>Y29468>YP4495
Q2b shew first up in ancient DNA in Sappali Tepe (Post-BMAC), but the sample was likely misdated and rather lived in the Iron Age than in the Bronze Age. He was of likely mixed Central Asian and West Iranic origin, but his Y-DNA was likely local or if from West Iran ultimately also derived from South Central Asia. So it is not surprising that in the Iron Age we see Q2b-Y1150 in regions in or close to the Inner Asian mountain corridor. 3 samples from the Swat valley in North Pakistan dated from 300 to 900 B.C were Q2b-1150. But it seems they were basal Q2b-Y1150 and negative for both downstream Q2b-YP755 and Q2b-Z5901. Two Loenbanr samples of higher coverage fall under the specific Q2b-BY186390 clade, which is a specific Q2b-Y1150 subbranch seemingly extinct or extremely rare today. Still considering the TMRCA of Q2b-Y1150 estimated to be around 3.500 B.C these samples are very interesting data points and probably derived their Q2b-Y1150 from a close IAMC population close related to the ancestor of Q2b-YP755 and Q2b-Z5901. These samples like many of the Iron Age Swat valley samples carried both Steppe MLBA and Inner Asian Mountain corridor admix. We also have Q2b-Z5901 downstream of Q2b-Y1150 from an Hunnic era sample from the Tian Shan region dated to 300 A.D but the sample was a genetic outliner with extra South-Central Asian ancestry. Today most Q2b-Y1150 in Afghans and neighbouring countries falls either under Q2b-YP755 or Q2b-Z5901.
Q2b-YP755 has a high frequency among KPK Pashtuns, Dehgan and Yusufzai and was also found among Pashtuns in eastern Afghanistan. With a TMRCA of around 5000 ybp and distribution from Xinjiang (Sarikoli, Wakhi) to Pakistan (mainly KPK) it likely is linked with some Bronze Age migrations of early Indo-Iranians. But a pre-Aryan migration into Afghanistan and North Pakistan can not be excluded for now. Q2b-YP755 was also found among Azeri and Poles so possibly was absorbed by Iron Age Steppe groups or Huns/Turkics later. Unlike other Q clades with a similar TMRCA found among Afghans and South Central Asians it seems to be absent in India.
Q2b-Z5901 with a TMRCA esimtated to be around 3400 B.C shows a wide distribution from South India to Tatarstan. It was also found among modern-day Hazara in Afghanistan and people of Khyber Pakhtunkhwa. Unlike Q2b-YP755 we already have an ancient sample under this clade. DA127 was Q-Z5901>YP4495 and orginated from the Hunnic era Tian Shan region but he likely had recent ancestry from the south. Based on the modern distribution and especially presence in South Indians it is likely that Q-Z5901 subclades like Q-Y28565 were picked up by early Indo-Aryans in the IAMC region and than further spread southwards in the Bronze Age. But it is likely that other Q2b-Z5901-rich groups remained in the Inner Asian Mountain corridor region up to the historic era and would be assimilated by Saka, Huns and Turco-Mongols.
Q-L717 in Pashtuns and South Asians
Q-L717 is a common Pashtun Q-L56 (Q1b) clade that was already found in the HGDP Pashtuns and Lodi Pashtuns. Its modern distribution from Kazakhstan/China to South India and TMRCA of 3400 ybp (Yfull) points strongly to a founder effect among early Indo-Iranians and especially Indo-Aryans of the Inner Asian Mountain corridor region carrying it further southwards in the Late Bronze Age and Iron Age. The closest ancient DNA relative to Q-L717 is Tutkaul 1 from Neolithic Tajikistan but it split more than 13000 years ago from the ancestor of Q-L717 so obviously not ancestral but points to the ancestor of Q-L717 populating the region between the Altai and Pamir mountains in the Neolithic and before.
This wide and diverse distribution of many Q clades makes it hard to pinpoint the exact origin of every Q clade in Afghans and surrounding people. We need both more modern and ancient samples to understand the migration route of Q clades. Most of the Q clades mentioned here often have very old TMRCAs and arrived probably not just in a single wave in Afghanistan.
BY186390 Loenbanr specific
Posth, Cosimo, et al. “Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers.” Nature 615.7950 (2023): 117-126.
Narasimhan, Vagheesh M., et al. “The formation of human populations in South and Central Asia.” Science 365.6457 (2019): eaat7487.
Raghavan, Maanasa, et al. “Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans.” Nature 505.7481 (2014): 87-91.
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